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Bulletin of the
British Museum (Natural History)
Entomology series Vol 37 1978
British Museum (Natural History)
London 1979
Dates of publication of the parts
No 1 : F : ; ; : : ; : : 25 May 1978
Nov yz : : : : : : : : ‘ 25 May 1978
No 3 : : ; : : 5 3 : : ; 25 May 1978
No4 . : : ; ; : ; : : : 27 July 1978
Nos: < . : : ; ; : : , : : 28 September 1978
NoG:. .. : ; : : ‘ ‘ : : : 30 November 1978
ISSN 0524-6431
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset
No 1
No 2
No 3
No 4
No 6
Contents
Entomology Volume 37
The anomalous ant-attended mealybugs (Homoptera: Pseudococcidae)
of south-east Asia.
D. J. Williams
A supplementary catalogue of the family-group and genus-group names
of the Coleophoridae (Lepidoptera).
K. Sattler & W. G. Tremewan
A catalogue of the type-specimens of the Scarabaeinae (Scarabaeidae)
and the smaller lamellicorn families (Coleoptera) described by G. J.
Arrow.
M. E. Bacchus
A revision of the Neotropical wasp genus Trigonopsis Perty (Hymen-
optera: Sphecidae).
C. R. Vardy
Nine genera of fungus-feeding Phlaeothripidae (Thysanoptera) from
the Oriental Region.
J. M. Palmer & L. A. Mound
The phlebotomine sandflies (Diptera: Psychodidae) of the Oriental
Region.
D. J. Lewis
Page
73
a7
18 Wy
153
PA
. —
/
> GENERAL
Bulletin of the | \, noe) |
British Museum (N: bial History)
The anomalous ant-attended mealybugs
(Homoptera: Pseudococcidae) of south-east
Asia
D. J. Williams
Entomology series Vol 37 Nol 25 May 1978
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology and Zoology, and an Historical series.
Parts are published at irregular intervals as they become ready. Volumes will contain about
four hundred pages, and will not necessarily be completed within one calendar year.
Subscription orders and enquiries about back issues should be sent to: Publications Sales,
British Museum (Natural History), Cromwell Road, London SW7 5BD, England.
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1978
ISSN 0524-6431 Entomology series
Vol 37 No 1 pp 1-72
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 May 1978
' 6 JUN j97¢
|
\y ,, LIBRARY ay
The anomalous ant-attended mealybugs \%;, 0. yy
(Homoptera: Pseudococcidae) of south-east “Asia
D. J. Williams
Commonwealth Institute of Entomology, c/o British Museum (Natural History), Cromwell Road,
London SW7 5BD
Contents
Synopsis . 1
Introduction , : : . i : ‘ : : : : 1
Habit. : : , é ; ‘ : : : ’ : : : 2
Distribution . : ; : : ‘ : : s : . ; 4
Morphology . ‘ ; : : : ; : : ; P : ; 6
Classification . ; “ : ‘ : : ; : ; @ : 4.9113
Acknowledgements . F : ; ‘ : : ; : : ; iy ts
Taxonomic treatment 3 : : : ; : : ; ; ; : 13
Pseudococcinae . , ‘ ; ‘ ; ; : ; F . 14
Allomyrmococcini trib. 1 Hie. f-; ; : ; i . 14
Key to genera of Allomyrmococcini (adult females) : : ‘ . ola
Key to genera of Allomyrmococcini (immature instars) . ‘ , : ; 15
Allomyrmococcus Takahashi, 1941 ; : : i , : : © er
Hippeococcus Reyne, 1954 . : : . : : : : : :. =20
Malaicoccus Takahashi, 1950 5 , , ; : : ; = 50
Paramyrmococcus Takahashi, 1941 ; ; ? ; ‘ : : . 54
Rhizoecinae . ; 62
Key to adult females of the anomalous ant- attended genera of Rhizoecinae ce Os
Eumyrmococcus Silvestri, 1926. , / ; ; : : , . § 63
Xenococcus Silvestri, 1924 . A ‘ ’ ; : : : ; is wis
References. : : ; 2 : : : ; = : : cog FID,
Index . ; : : 5 ; ; ; : : ; ¢ ‘ yf
Synopsis
The morphology, habits and taxonomy of 12 unusual myrmecophilous mealybugs from south-east Asia
are discussed. Unusual characters of some of the species are long antennae, an anal ring without pores,
a dense covering of minute setae on the dorsum, long anal lobe setae and large protruding ostioles. The
ostioles are unlike any others in the Pseudococcidae and are compared with the siphunculi of aphids.
Allomyrmococcus, Hippeococcus, Malaicoccus and Paramyrmococcus are aerial genera and are assigned
to the subfamily Pseudococcinae. These four genera have many characters in common and a new tribe
Allomyrmococcini is erected for them. Two new species of Malaicoccus and one new species of Para-
myrmococcus are described. Eumyrmococcus and Xenococcus are subterranean and are placed in the sub-
family Rhizoecinae.
Introduction
Mealybugs are easy to recognize in life by an elongate-oval body usually covered with mealy or
cottony white wax. This wax often extends around the sides to form a series of short filaments
but a pair of much longer filaments may be present at the posterior end of the body. Most species
have legs which rarely protrude beyond the lateral margins and movement is usually slow.
Oviparous females become sedentary and may secrete a noticeable white cottony ovisac.
Bull. Br. Mus. nat. Hist. (Ent.), 37 (1): 1-72 Issued 25 May 1978
In common with all female scale insects, species can be identified only from microscope slide
mounts, by the arrangements of body pores and setae. Mealybugs are separated from related
groups in possessing at least one of the major characters, cerarii, ostioles, circulus, trilocular
pores and tubular ducts which are not cupped at the interior end.
As in all Homoptera, scale insects suck the sap of plants with long stylets and, so far as is
known, all mealybug excretion is in the form of honeydew. This substance is excreted in all the
families of Coccoidea except the Diaspididae, Conchaspididae, Halimococcidae and certain
groups at present assigned to the family Asterolecaniidae. When palatable, honeydew forms an
important part of ants’ diet and a strong association has been built up between some species of
ants and certain Homoptera. The literature discussing this association is extensive and in recent
years Nixon (1951), Way (1963) and Wilson (1971) have discussed the subject at some length.
Way has defined mutualism as any association beneficial to both the ant and the other insect.
Myrmecophilous insects (according to Way) are those which benefit from ants and are more or
less adapted to live with them but the relationship need not be obligatory or mutually beneficial.
The majority of myrmecophilous mealybug species having a mutual association with ants
are those which a coccidologist would regard as normal and without visible signs of special
adaptation. In south-east Asia some myrmecophilous mealybugs have developed most unusual
characters and habits and it is the purpose here to describe and discuss this group. At present
the group comprises the genera Allomyrmococcus Takahashi, Eumyrmococcus Silvestri, Hippeo-
coccus Reyne, Malaicoccus Takahashi, Paramyrmococcus Takahashi and Xenococcus Silvestri.
Habit
Silvestri (1924) described the first of these mealybugs under the name Xenococcus annandalei.
It is an elongate-oval species but the abdomen tapers abruptly at the posterior end, and the long
antennae are strongly geniculate with a special articulatory mechanism between the first and
second segments. One of the most noticeable characters is the dense covering of minute setae
on the dorsum which at the time of description had not been seen in mealybugs before. This is
a subterranean species found in the nests of the ant Acropyga acutiventris Roger. Silvestri
described the habit as living on the rootlets of Ficus sp. When the soil is damp and warm both
ants and mealybugs remain just below the surface under stones but in cold, dry weather they
retire deep into the ground. The ants carry away mealybugs in their mandibles when the nest is
disturbed and when ants leave the nest, each female carries a female mealybug in her mandibles
to new nests elsewhere.
Later Silvestri (1926) described Eumyrmococcus smithii from Macao and Shanghai as living
under stones with the ant Acropyga (Rhizomyrma) sauteri Forel. Recorded later in Taiwan with
the same ant, Takahashi (1934) mentioned that females and workers carry the mealybugs in the
mandibles. This species is also densely covered with minute setae and the abdomen is abruptly
narrowed but the antennae are short.
Above ground in Thailand were found Allomyrmococcus acariformis and Paramyrmococcus
chiengraiensis described by Takahashi (1941). Both were covered with dense clusters of the ant
Dolichoderus sp. [=Hypoclinea]. Malaicoccus riouwensis described by Takahashi (1950) from the
Riouw Is (now known as Riau Is) and M. formicarii described by Takahashi (1951) from Malaya
were both attended by large numbers of Polyrhachis sp. This ant carried M. formicarii in its
mandibles at the constricted part of the thorax. A new species described here as M. moundi
is also carried at the head or prothorax by the ant Hypoclinea sp. (Fig. 1B). Specimens of the
mealybug examined in alcohol were firmly grasped in the ants’ mandibles which had to be prised
apart to release the mealybugs.
Species of Hippeococcus described by Reyne (1954) have an even more unusual habit. These
species are also aerial, feeding on stems and suckers of various plants, and are covered with ants
of the genus Dolichoderus (= Hypoclinea). Mealybugs of H. rappardi Reyne are gleaming without
any trace of wax and are very agile. When the ants are disturbed the mealybugs cling to the body,
sitting crosswise on the ants’ back (Fig. 1A). Immature H. wegneri Reyne feeding on young shoots
and fruit of Diospyros kaki cling to the thorax and other parts of the ant’s body. A Hypoclinea
y
Fig.1 (A) Ant, Hypoclinea sp., carrying Hippeococcus sp. on its back. Redrawn from Reyne (1954).
(B) Ant, Hypoclinea sp., carrying Malaicoccus khooi in its mandibles. Drawn specially for this work
by Mrs Linda Huddleston of the British Museum (Natural History).
3
nest is also aerial and sometimes contains numerous mealybugs which, according to Reyne,
cling to the ants’ bodies when the nest is opened. Ants also drag along swollen adult females and
sometimes carry mealybugs in their mandibles. Reyne mentioned that ants were often seen drum-
ming incessantly on a certain spot on the mealybugs and although honeydew was produced as a
clear drop at the tip of the abdomen, the ants showed little interest in it. When H. wegneri,
living on Cyclanthera explodens, was disturbed, the ant colony assumed an alarm position and
the ants evacuated the mealybugs by first tapping each mealybug with the antennae and then
carrying the mealybugs on their backs. Larger mealybugs were often given preference and after
the disturbance the ants returned to unload the mealybugs.
These observations reveal an unusual and possibly old association between ant and mealybug
but leave many questions unanswered. Broadbent (1951) stated that aphids excreted honeydew
only when feeding and if mealybugs excrete in the same manner, it is strange that ants were never
seen to take honeydew from Hippeococcus on the stems. There were no observations as to whether
the mealybugs were feeding inside the nests or whether ants were soliciting them. It is difficult
to imagine such a close relationship without ants receiving honeydew at some time and it must
be assumed that they take it. No sooty moulds were observed growing on any honeydew falling
on plant material. Way (1954) reported that when the ant Oecophylla longinoda (Latreille) in
Zanzibar was removed from the Coccid Saissetia zanzibarensis Williams there was a noticeable
fall in the Coccid population because of increased parasitism and suffocation from the Coccid’s
own honeydew. Das (1959) reported similar suffocation of Coccus hesperidum L. on tea in
Assam when ants were excluded. The honeydew apparently oozes slowly rather than being
ejected for some distance as in Pulvinaria ribesiae (Signoret) reported by Newstead (1903).
Furthermore, during periods of heavy rainfall in Zanzibar, O. longinoda workers were often
unable to remain in attendance on the Coccid and the honeydew was washed on to the surround-
leaves. Hippeococcus and Malaicoccus live in areas where heavy rain beats on ant and mealybug
for long periods without any lessening of ant activity. Biinzli (1935) reported that the ant Rhizo-
myrma paramariensis in Surinam carried Rhizoecus [ =Neorhizoecus] coffeae Laing and Geococcus
coffeae Green from wetter to drier areas in underground nests. Way (1954) noted that Oecophylla
longinoda kept Saissetia zanzibarensis numbers at a level which satisfied the honeydew require-
ments. Flanders (1957), discussing the obligate myrmecophilous mealybug Chavesia sp. and the
obligate coccidophile Rhizomyrma fuhrmanni Forel, stated that the mealybug can feed only on
roots exposed by the ant. Furthermore, the worker ant regulates the number of mealybugs feeding
at any one time by placing about 70 per cent of the total mealybug population in reserve in a
non-feeding position off the roots. How ants remove mealybugs from stems without damaging the
stylets has never been observed. In Hippeococcus there are apparently numbers of mealybugs on
the stems and even greater numbers in ants’ nests and one can only speculate that this is a form
of regulation by the ant. Hypoclinea is not an obligate coccidophile and the question remains
whether Hippeococcus and the related Malaicoccus, Allomyrmococcus and Paramyrmococcus
are obligate myrmecophiles. The answers probably lie in the unusual characters of these mealy-
bugs and these are discussed in the section on morphology.
Distribution
Although Silvestri described Xenococcus annandalei from Lake Chilka near the West Bengal
border, it has been discovered since in Mysore, southern India and Penang in West Malaysia,
northern Vietnam and Hong Kong, always associated with the ant Acropyga acutiventris Roger.
Eumyrmococcus smithii is known from Macao and Shanghai in China and in Japan and Taiwan
associated with the ant Acropyga sauteri. These two mealybugs are subterranean and probably
feed on a wide variety of roots.
Among the aerial species, Hippeococcus is known from Java only. Malaicoccus is known from
Malaya and the Riouw Is in Indonesia but based on the identity of immature stages in this work,
a third stage of an apparently new species was mixed with Allomyrmococcus acariformis when
Takahashi described it from Thailand. Paramyrmococcus is discussed here from Thailand and
4
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Vietnam. These aerial species seem to be attended only by ants of the genera Hypoclinea and
Polyrhachis which are widespread in south-east Asia. The mealybugs will probably have a distri-
bution throughout the area wherever the ants are present. Since this paper went to press the
writer has received a new species of Malaicoccus attended by Hypoclinea sp. from Sarawak near
the border with Sabah.
Morphology
Body
In life the body shape tends to be clavate with the head and thorax dilated and with the abdomen
narrowing abruptly at about the third segment. Only in Malaicoccus is the abdomen broadly
rounded. The body of Xenococcus is widest at about the first abdominal segment but the body
still has a clavate appearance by the enormous basal segments to the antennae. Paramyrmococcus
is more turbinate. In South America Chavesia has a similar body shape to that of Eumyrmococcus.
Chavesia seems to have developed along similar lines to the genera in south-east Asia and there
is strong evidence that Xenococcus and Eumyrmococcus are closely related to Chavesia but that
Allomyrmococcus, Paramyrmococcus, Malaicoccus and Hippeococcus form a separate group.
Coccidologists become used to studying flattened insects on slides, discussing only dorsal and
ventral surfaces and forgetting that species have lateral sides. In all the genera from south-east
Asia and South America the abdomen curves upwards at the posterior end, as shown by Silvestri
(1924; 1926) and by Balachowsky (1957). It is not clear what advantage this gives to the mealybug
or the ant. When an ant strokes a mealybug the abruptly narrowed abdomen may resemble an
ant petiole and so the mealybug may be more readily accepted. Ants seem to carry these mealy-
bugs near the head and prothorax whereas ants carry mealybugs of more normal shape at about
the first abdominal segment. As Biinzli (1935) has remarked, the mandible marks may disappear
but sometimes they may persist. Malaicoccus has developed heavy sclerotization on the head at
the point where they are carried.
Legs
The legs of all species are well developed and reach their greatest length in Allomyrmococcus
when they are longer than the body. Reyne (1954) has mentioned that Hippeococcus moves
quickly yet Strickland (1950) stated that in West Africa the mealybugs associated with ants
have shorter legs than the active longer legged species that are not associated with ants. The leg
setae are well developed on each species and sometimes they are abundant. The claw gives some
clue that these mealybugs belong to two distinct groups. In Xenococcus and Eumyrmococcus
the claw is long and slender with a pair of short setose digitules at the base but in the other four
genera the claw is stout with a pair of large flattened digitules which are usually twice as long as
the claw. Reyne (1954) has suggested that Hippeococcus uses the digitules to cling to the ants’
bodies and this seems a reasonable deduction. It should be remembered that these large flattened
digitules are present in Allomyrmococcus, Paramyrmococcus and Malaicoccus also but these
mealybugs apparently do not cling to ants’ bodies. The tarsus of each species has a single cam-
paniform sensillum at the outer proximal end, a structure discussed recently by Koteja (19745).
Antennae
Eumyrmococcus has short 2-segmented antennae but in Xenococcus the antennae are 4-segmented
of a most unusual type. These are long and strongly geniculate with a large basal segment. The
articulation between the first and second segments is so well developed that the antennae can
bend from a forward position to one lying along the back of the mealybug. Helping in this move-
ment are small teeth on the outer proximal corner of the second segment which fit into grooves
on the outer distal corner of the first segment. Long pointed setae are present on all the segments
and on the third and fourth segments some setae are as long as the segments. As Xenococcus
lives underground it is probably advantageous for the mealybug to streamline its body by folding
6
the antennae along the back but in other circumstances the antennae may be used for recognizing
an ant.
The antennae of the other four genera are 6-segmented and are unlike those of any other mealy-
bugs. In each species and in all stages, the second segment is short and does not articulate with
the third segment. In the first stage larvae, long setae form whorls mainly at the distal ends of
the segments but in Paramyrmococcus vietnamensis the fifth and sixth segments have numerous
short setae also. Short setae are present on all second instars and they become more numerous
and longer on each successive stage so that the adult antennae are densely covered. The antennae
in most adult females of this group are about as long as the body. It is difficult to understand
why these insects have such long and well-developed antennae. It is usual for an ant to feel or
stroke a mealybug before either attending one or carrying one away. Mealybugs of Hippeococcus
and possibly those of related genera may equally seek out or recognize an ant with their long
antennae.
Setae
One of the striking characters of nearly all these mealybugs and the South American genus
Chavesia is the dense covering of minute setae over the entire body or at least over the dorsum.
In many species the setae are short and are so numerous that the distance between the setal bases
is much shorter than the length of a single seta. These mealybugs are without the usual protection
of mealy or flocculent wax and the numerous setae may act as an alternative protective covering.
By trapping air between the setae, the body surface of underground species would be protected
from excessive moisture. The aerial species are exposed to heavy rains for long periods and the
dense covering of setae may have a waterproofing effect.
Some species do not have such dense coverings. Adult Hippeococcus have, in addition to minute
pointed setae, small blunt setae of uncertain function. Longer clavate and fleshy setae, which in
H. rappardi are present in the first stage only, persist in all stages in H. wegneri. In many of the
immature stages of Malaicoccus there are minute setae in varying numbers, each with the seta
shorter than the diameter of the setal base. Immature Paramyrmococcus have body setae which
are short, cylindrical and blunt but these are replaced by numerous short pointed setae in the
adult. A few setae which are flat and variously shaped are present on the head and thorax of
Xenococcus. There is little difference, apart from abundance, in the arrangement of the setae in
the stages of Allomyrmococcus but immature stages have extra long setae on parts of the body.
Adult mealybugs, therefore, usually have some characters such as the addition of special body
setae or the loss of long setae, which distinguish them from the immature stages. Ants may be
able to recognize these differences by quickly stroking the mealybug’s body.
Anal areas
This section includes the anal ring, anal lobes, anal lobe setae and any other structures at the
apex of the abdomen. The anal ring is usually sclerotized with 6 setae of various lengths and always
without pores except for a few minute pore-like openings in Malaicoccus. In the more usual
mealybugs, of which Pseudococcus obscurus Essig is an example, a short wax tube is secreted
from pores around the anal ring. Honeydew droplets are, therefore, excreted a short distance
from the body surface. A colony of P. obscurus on a potato tuber at times ejected honeydew for
a distance of 2-4 cm. This honeydew is clear and sticks to the surrounding foliage, but each droplet
of honeydew of the cochineal insect Dactylopius coccus Costa is coated with fine wax and falls
directly to the ground without sticking to the plant. Broadbent (1951) has mentioned that the
aphid Hayhurstia atriplicis (L.) excreted globules coated with wax from contact with the wax
covering of the aphid. There is always the possibility that wax coating may affect the palatability
to ants. No observations have yet been made on the honeydew of the myrmecophilous mealybugs.
Reyne (1954) has stated that Hippeococcus rappardi had usually a clear drop of liquid between
widely diverging silvery hairs, which are presumably the anal hairs. In Chavesia, Beardsley (1970)
has described the anal lobes as protuberant and sclerotized on the ventral surface. The anal ring
is in a membranous depression on the dorsal surface of the lobes. These lobes also contain
7
numerous setae and the whole structure forms what Beardsley has called a honeydew basket. A
similar structure is present in A/lomyrmococcus and Malaicoccus but the anal ring may be ter-
minal or situated a short distance from the apex. In Hippeococcus and Xenococcus the anal ring
lies at the apex of the dorsum which projects beyond the ventral anal lobes. Wherever the anal
ring is placed it is within easy access for the ant and in the absence of pores it seems likely that
honeydew remains at the apex of the abdomen until the ant removes it. Broadbent (1951) has
described how the aphid Hyalopterus pruni (Geoffroy) kicks away globules with the hind legs.
Normally the legs of mealybugs are too short to reach the anus but in the absence of ants most
of the myrmecophilous mealybugs in this discussion could remove globules with their legs if
necessary but this has not been observed.
Long anal lobe setae are present in all genera except Paramyrmococcus. Sometimes they are
numerous and are longer than the body. Their function is uncertain and their absence in Para-
myrmococcus does not seem to affect the myrmecophilous habit.
As previously stated, honeydew is probably produced only when the mealybugs are feeding.
Furthermore excretion rates may be increased by ants attending mealybugs and removing honey-
dew. It is usual for an ant to stimulate the mealybug into giving up a drop of honeydew by pal-
pating the abdomen, but the mechanism of this stimulation is not understood. In the Coccidae,
Way (1954) and Smith (1942) have discussed how ants solicit species of Saissetia by dragging
the tips of the antennae back and forth over the caudal end of the body. Saissetia and Udinia are
two related genera often solicited by ants in the tropics but species of both genera have long pri-
mary discal setae on the anal plates which when touched may induce the Coccid to open the plates
and excrete some honeydew. By stroking the anal ring setae, ants may induce mealybugs to excrete
a droplet.
Body pores
An important feature of the myrmecophilous mealybugs is the almost complete absence of pores
and ducts. All species have been described without surface wax, and Reyne (1954) described
Hippeococcus as having a gleaming body. Eumyrmococcus and Xenococcus are completely without
body wax pores and in all the other genera the only pores present in the female stages are trilocular
and a few simple pores except in some species of Malaicoccus which have a few oral collar ducts
in the mid-thorax and anterior abdominal segments. In most species the trilocular pores are quite
sparse but in Hippeococcus and certain Malaicoccus species they are more numerous. It is only
the second stage male which has the more usual pores associated with mealybugs. In H. wegneri
and M. moundi the second stage males have multilocular disc pores and oral collar tubular ducts
and it is assumed that these secrete the waxy covering for the pupal stage. Nixon (1951) has given
a few examples of ants removing wax coverings of mealybugs. For instance, Le Pelley (1943)
noted that ants removed wax completely from certain areas of the body of Pseudococcus [ =Plano-
coccus] lilacinus (Cockerell) in the Philippines. It is not clear how male mealybug pupae could
retain wax coverings in the presence of attending ants. Nevertheless, the gleaming and waxless
condition of myrmecophilous mealybugs is probably the result of specialization in the continual
presence of ants. Recently Delage-Darchen et al. (1972) have noted an unusual association between
Diaspidid scale insects and ants. The scale insects live in tunnels in woody stems and are attended
by ants in so far as ants remove the scale coverings. One species of these scale insects is completely
without dorsal ducts so that no scale could be secreted anyway. Similar observations with scale
insects have also been made by Prins et al. (1975). Although ants may remove wax for food it is
also possible that in time species that would normally secrete a wax covering evolve without even
any means of producing one. It is difficult to understand how ants benefit from species not producing
honeydew.
Labium
The labium of Pseudococcidae is conical and 3-segmented and its morphology has been studied
by Koteja (1974a) whose terminology is used here. The basal segment is small with a pair of
rounded sclerites, often fused medially with the medial segment, and contains 2 pairs of short
8
setae and one longer pair. On the medial segment there is a basic number of | pair of setae. The
apical segment has 2 or 3 pairs of posterior setae on the posterior or under surface, a pair of short
apical setae, 4 pairs of subapical setae, one pair of lateral setae and 2 pairs of anterior setae
situated near the base. In many species there are supernumerary setae anterior to the lateral setae.
Koteja recognized the four subfamilies Trabutininae, Pseudococcinae, Rhizoecinae and
Sphaerococcinae. The Trabutininae is separated from the others, using the character of the
labium only, in possessing 3 pairs of posterior setae. All the species studied in this work have 2
pairs of posterior setae and the species are separated from the Sphaerococcinae in having a well-
developed basal segment not fused totally with the medial segment. The labium of Rhizoecinae,
according to Koteja, differs from that of the Pseudococcinae in being small and narrow with the
third subapical setae distinctly stouter than the others and with the lateral setae about twice as
long as the other labial setae. Williams (1969) has already placed Eumyrmococcus in the Rhizoe-
cinae on other grounds but the labium of this genus has a width-length ratio of 1: 2, there are
14 pairs of setae and the subapical setae become progressively longer towards the base. Xeno-
coccus has a labial width-length ratio of | : 1-87 when flattened on the slide and | : 2-27 when not
flattened. The number of setae varies from 12 to 13 and the subapical setae are remarkable in
extending more than half way to the base of the apical segment. As in Eumyrmococcus these
subapical setae become progressively longer towards the base of the segment. For reasons to
be given later under classification, Xenococcus should be assigned also to the Rhizoecinae.
In the genera Hippeococcus, Malaicoccus and Paramyrmococcus the width-length ratio of the
labium varies considerably from 1: 1-4 in P. chiengraiensis to 1: 1:92 in Malaicoccus khooi.
Despite this range in size there is reason to believe that these genera with Allomyrmococcus
belong to the Pseudococcinae. In Allomyrmococcus and Malaicoccus the total number of setae
varies from 20 to 32 pairs in the adult female but in the first and second stages they become
reduced to the basic number of 14 pairs.
Circulus
The circulus, when present, is a round or oval sclerotized area, surrounded by a rim, usually
lying on venter between the fourth and fifth abdominal segments. Occasionally it may lie within
the borders of the fourth segment and some species may have several circuli on different segments.
In the Rhizoecinae the circulus is usually small and round but heavily sclerotized, conical and
often with an irregular surface. Pesson (1939a) and Lloyd & Martini (1957) have shown that
adult females of the genera Pseudococcus or Phenacoccus use the circulus as an adhesive organ
when viewed in profile against a glass surface. The circulus is exserted and then adpressed to the
surface allowing the insect to make exploratory leg movements, the body actually pivoting at the
circulus. This use seems to be restricted to the adult female because Lloyd & Martini observed
that the three nymphal instars were able to ascend or descend with ease by leg actions presumably
with the help of the claw digitules.
No similar research has been done on the circulus of the Rhizoecinae but Silvestri (1924),
working on alcohol material of Xenococcus annandalei, suggested that the circulus of this species
secreted some kind of liquid possibly attractive to ants. Whatever its function it does seem that
the circulus of the Rhizoecinae has been derived from those of the other groups of the Pseudo-
coccidae.
Ostioles
Among the Coccoidea the Pseudococcidae are unique in usually possessing 2 pairs of transverse
slit-like organs on the dorsal surface, commonly named ostioles. In Allomyrmococcus, Hippeo-
coccus, Malaicoccus and Paramyrmococcus the ostioles reach an unusual development and this
seems an appropriate place to discuss their structure and to compare them with siphunculi or
cornicles in aphids.
Occasionally some species of such genera as Rhodania Goux and Ehrhornia Ferris are without
ostioles but they may be present in some of the immature stages where they are limited to the
posterior pair only. In adult females of Antonina, ostioles may be present as a poorly developed
9
posterior pair only. The absence or reduction of ostioles is correlated with the amount of external
wax secreted from the pores, the term wax being used here in its general sense. Usually those
species producing large amounts of dorsal wax or those which become almost enclosed in a felted
Ovisac, as in Antonina, have reduced ostioles.
According to Ferris & Murdock (1936) the anterior pair belong to the pronotum and the
posterior pair to the sixth abdominal segment. Pesson (1939b), however, after an overall study of
the musculature, concluded that the anterior pair are placed slightly forward to the anterior edge
of the prothorax and the posterior pair lie between the fifth and sixth abdominal segments.
Viewed from above each ostiole is composed of two membranous lips which often contain
numerous setae and trilocular pores. The inner edges of the lips are usually sclerotized and some-
times these inner edges are the only parts visible.
Sulc (1909) observed in Phenacoccus farinosus (de Geer) that blood corpuscles oozed from the
ostioles after pressure was applied to the body. He also observed some muscles parallel to and
perpendicular to the lips of the ostioles.
In histological studies Ferris & Murdock (1936) found no muscle attachments on these struc-
tures and concluded that they led directly into the haemocoele. When the insect was stimulated
by an application of violence, a globule of liquid appeared which quickly hardened on exposure
to air. The globule, according to Ferris & Murdock, was merely a portion of the body fluids.
In more detailed histological studies, Pesson (1939b) found that in a transverse section, the edges
of the slit are continued into the body cavity as a chitinized depression, the median part of which
forms an orifice leading directly to the haemocoele. From the lateral inner ends of the depression
in each anterior ostiole are inserted two muscles, the anterior or inner muscle inserting itself
obliquely to the inner edge of and lateral to the clypeus, and the posterior or outer muscle
attached a little anterior to the first leg. There are also two similar muscles in each posterior
ostiole which are attached ventrally. If an ostiole is regarded as a valve then normally it should
prevent blood loss when at rest but after an application of some body violence near an ostiole
the muscles contract and open the lips slightly to emit a droplet of blood or other contents of the
haemocoele. This emission, according to Pesson, is nothing more than reflex-bleeding or auto-
haemorrhage.
Sulc (1909) noted that the expelled wax, on hardening in contact with air, could have a protect-
ing roll by coating the oral parts of an attacker. Pesson never witnessed this in Pseudococcus
adonidum (L.) [=P. longispinus (Targioni-Tozzetti)] and indicated that the liquid expelled is
from the free wax occurring in the blood as corpuscles. After repeated excitation of the surround-
ing cuticle young wax-producing cells may be emitted. Wheeler (1921), in his studies of the myrme-
cophyte Tachigalia and the attending behaviour of beetles of the genus Coccidotropus on Pseudo-
coccus bromeliae (Bouché) [= Dysmicoccus brevipes (Cockerell)], observed that the beetles were
interested only in the honeydew excreted and were never immobilized or impeded by any secretion
from the ostioles. He stated that the beetles may be attracted by some fascinating aroma secreted
by the mealybug but did not actually state that this was emitted from the ostioles.
After Coccoidea are macerated in potash as a first step in the mounting technique there are
often observed certain internal wax globules varying in quantity as to the family. It is in the
Pseudococcidae that this internal wax is most strikingly abundant and it is apparently this free
wax which is expelled through the ostioles.
The posterior ostioles have often been homologized with the cornicles or siphunculi of aphids.
Little work has been done so far on ostioles but in recent years work on siphunculi has been
extensive. A résumé of recent work on siphunculi in aphids is given here and it is hoped this may
stimulate further work on ostioles.
It was shown by Hottes (1928) that cornicles are situated on the dorsolateral surfaces of the
sixth abdominal segment. In their most conspicuous form they are cylindrical and longer than
wide. Usually the longer cornicles are movable but at the tip of all cornicles there is a slit-like
opening which may be opened or closed by a valve moved by a muscle attached near the median-
free portion of the valve. The valve is attached at the apex by a flexible hinge. Hottes further
indicated that, although cornicles may be movable, the possibility that droplets can be aimed is
practically nil. The function of the cornicles, according to Hottes, was not a defence mechanism
10
but was part of the metabolic processes carried out by the aphid and that the cornicles have an
excretionary function. Furthermore, the presence or absence of cornicles may be correlated
with the presence or absence of wax pores or a lower reproduction rate. Hottes stated that as
aphids have often evolved along with their hosts the change in structure of the cornicles may be
the result of the nature of the food consumed.
Despite Hottes’ rejection of the defence mechanism function of cornicles, Dixon (1958)
observed that if the larva of the coccinelid Adalia decempunctata (L.) was smaller than the aphid
Microsiphum evansi (Theobald) and seized an appendage, the aphid could escape by pulling the
appendage free. If the coccinelid larva and aphid were about the same size the aphid could escape
if the siphunculus nearest the appendage swung over and placed a drop of oily liquid on the
coccinelid’s head where the drop would spread over the mouthparts and solidify.
Reflex-bleeding discussed by Hottes was confirmed in aphids by Edwards (1966) who stated
that when the cornicle valve is opened, the aphid released material from the haemocoele. As no
solvent of the internal material was detected by gas chromatography he explained that the fluid
is in a stable liquid—crystalline state within the aphid and changes to the solid crystal phase with
a seeding nucleus. He further stated that the rapid crystallization on contact with a solid surface,
a hair or duct, suggests that the liquid was in a supercooled state and that foreign material
provides a seeding nucleus. The melting point of the waxes of three species of aphids ranged
from 37-5 °C to 48 °C so that the waxes should crystallize on seeding at normal summer tempera-
tures. Studying the cornicle area of various species of aphids, Wynn & Boudreaux (1972) found
that in Cinara and Longistigma a multicellular sac possessing a wall, formed from a layer of flat
cells with compressed nuclei, extends into the cornicle. This sac is surrounded by vacuolated fat
cells and a mass of lipid is found in the sac. All the aphids studied possessed a muscle inserting on
the cornicle valve and originating on the venter below the cornicle. The authors suggest that this
muscle, in addition to opening the valve at the distal end of the cornicle, may also assist in elevat-
ing the cornicle. They also suggest that a dorsoventral muscle just anterior to the cornicle appears
to be used in association with other dorsoventral muscles in compressing the body contents
causing the emission of a droplet. In each case the cornicle communicates with the haemolymph
and blood cells including fat cells escape. The colour of the cornicle droplet corresponds with the
colour of the aphid and its haemolymph.
In contrast, Lindsay (1969), studying the cornicles of the pea aphid Acyrtosiphon pisum (Harris),
stated that the term reflex-bleeding as applied to cornicle exudation is inapplicable since, although
a small amount of haemolymph may escape, the primary exudate is the cornicle cells. Analysis
of these cornicle cells indicates a close similarity to those of the fat-body cells in composition.
Lindsay suggested that the fat-body cells and cornicle cells are homologous, the fat body being a
precursor of the cornicle cells and that there is a greater tendency of the cornicle cells to crystallize
compared with those of the fat-body cells.
Evidence that haemolymph is not extruded through the cornicles was presented by Strong
(1967) who stated that lipids were the principal constituents of the hardened cornicle droplets
and were composed solely of triglycerides. Histological preparations by Chen & Edwards (1972)
showed that the cornicle secretionary cells are present inside a sac separated by a thin acellular
lamina from the haemolymph. The sac is formed by an invagination of the epidermal basal
lamina reflected back from the tip of the cornicle. The enclosed secretionary cells are comparable
with epidermal gland cells or subepidermal oenocytes. Fluid in the sac differs in composition from
the haemolymph and the fluids released through the cornicle are thus not a direct loss of haemo-
lymph but are presumably derived from the haemolymph.
Referring to the triglycerides in the cornicle secretions, Callow et al. (1973) found that the
secretions in any one species are the same and that different species of aphids taken from the
same host plant have different secretions. The authors found it was easier to obtain droplets
from young or small aphids rather than large aphids and they presumed that the larger aphids
have less need of a defence mechanism. This is interesting because as already stated ostioles may
be present in immature mealybugs and absent in adults although defence mechanisms in mealy-
bugs have not been proved there may be a greater need for ostioles in immatures for different
reasons. Analysis by Greenway & Griffiths (1973) of the body triglycerides showed them to contain
11
the same fatty acid radicles as in the cornicles but in many species there are greater proportions
of some triglycerides in the body. When an aphid is overstimulated the composition of the cor-
nicle secretions gradually becomes the same as the body contents.
Speculative suggestions that odours may be emitted by mealybug ostioles have not so far been
confirmed but Dahl (1971) reported that odours from crushed aphids repelled other apids. Kislow
& Edwards (1972) found that certain aphids are repelled by the odour from cornicle droplets
and squashed specimens of the same species. These authors proposed that when there is a slow
release of odour it may cause intraspecific spacing of aphids, prevent the influx of other species
or even induce aphids to migrate. The alarm pheromone was identified as trans-B-farnesene or
TBF in some aphids by Bowers et al. (1972). Nault et al. (1973) showed that the alarm pheromone
is known to be interspecific and that the receptor of this repellent odour is the primary sensorium
on segment 6 of the antenna. The existence of multiple alarm pheromones was suggested by Nault
& Bowers (1974) who found that Hydaphis erysimi (Kaltenbach) does not respond to pure TBF
and they suggest that the aphid produces one or more other seh in addition to TBF to
produce alarm activity.
Before referring once more to ostioles in mealybugs it is interesting that Dixon & Stewart
(1975) after blocking the openings of siphunculi of the sycamore aphid Drepanosiphum platanoides
(Schrank) found that this occlusion had no adverse effect on the, number of dffspring produced
or on the weight of the aphid. Furthermore, aphids with: occluded siphunculf do not space out
differently from aphids with normal siphunculi and that'the aphid takes most avoiding action
when the smell of the pheromone is accompanied by vibrations associated with a struggling aphid.
There is obviously a need for -much further research on ostioleés: -Eyen a cursory glance at the
ostioles of Allomyrmococcus, Paramyrmococcus and Hippeococcus shows that they are much more
highly developed than in normal mealybugs. The sclerotized lips have their greatest development
in any mealybugs known to the author. Although i in the iHustrations they are located on the edges,
this is due to dorsoventral flattening when specimens are prepared on microscope slides but even
in specimens preserved in alcohol they are unusually large and conspicuous with the sclerotized
lips protruding on the dorsolateral areas of the body. Their most unusual development is found in
Malaicoccus. In this genus the anterior pair are often located on the ventral surface on slide
preparations. Each ostiole appears to have a fixed anterior sclerotized lip but the posterior lip
is a large hinged semicircular flap often containing numerous setae. This flap is often seen com-
pletely closed but in some specimens it is open and raised at a right-angle.
In the four genera there is in each ostiole a membrane, easily taking up stain, which completely
covers the opening of the lips. It is not certain, working only from microscope preparations,
whether there is any opening on this membrane. There is in some specimens a hole or slit on the
surface but this may be due to tearing during the mounting technique. The membrane may nor-
mally be internal and may be exserted as a result of pressure on the body when flattened.
Why these species should possess such complicated ostioles is still uncertain but the mealybugs,
as explained earlier, are transported when the ants are disturbed. Although ants are attracted to
excretions of honeydew it is doubtful if honeydew has any odour over long distances but any
odour from the ostioles would attract ants and thus help them to transport mealybugs back to the
preferred host plants. This is conjecture but any work on pheromones must be done in areas
where the mealybugs are common. It is easy to bring aphids into the laboratory for study but
much more difficult to experiment with the most interesting of mealybugs which inhabit the
tropics. If there is an odour from ostioles of these myrmecophilous mealybugs to attract ants,
then there is possibly a similar odour emitted, as suggested by Wheeler, from those mealybugs
with a less obligatory association with ants. Any research on this odour could lead to an alteration
of the relationship between ants and mealybugs and could possibly help in controlling mealybugs.
Mutual association with ants in the Palaearctic Region is common in such mealybugs as Chnauro-
coccus subterraneus (Newstead) and Euripersia europaea (Newstead), species which may be useful
for research, and in the sugar-cane areas mealybugs living in association with ants are common
enough to afford the means for easy research. It is doubtful, for instance, whether alarm phero-
mones are emitted by mealybugs as in aphids but any work on mealybug ostioles that may
determine if their function is different from reflex-bleeding usually propounded would be useful.
12
Classification
Xenococcus and Eumyrmococcus seem to belong to the subfamily Rhizoecinae. Williams (1969)
placed Eumyrmococcus in this group because it possesses long slender claws with setose digitules
and 3 pairs of long anal lobe setae. Xenococcus has similar long claws and also 2 circuli which
are round, sclerotized and slightly conical. The antennae are not typical of the Rhizoecinae but
the reduction in the number of segments suggests a relationship. The long slender labium of both
genera each have a width-—length ratio of 1 : 2 and the assignment of the genera to the Rhizoecinae
agrees with research of the group made by Koteja (1974a; 1974c). Beardsley (1970) tentatively
placed the genus Chavesia in the Rhizoecinae and this is probably correct. Some of these species
have 4-segmented antennae but of a different shape to those of Xenococcus. The labium of
Chavesia has a width-length ratio varying from 1 : 1-33 to 1 : 2:00 but Beardsley has described it
as being 2-segmented only so that the addition of the basal segment would alter the width—length
ratio. The labium of C. trinidadensis Beardsley has a similar shape to the labium of Xenococcus.
The four genera Allomyrmococcus, Hippeococcus, Malaicoccus and Paramyrmococcus have
many characters in common. The antennae and claws are remarkably similar and so unusual
that a new tribe is here erected for the genera within the subfamily Pseudococcinae. In the absence
of adult males it is not possible to give the relationship of the tribe.
Separation of the subfamilies Pseudococcinae and Rhizoecinae in this work is based mainly
on the shape of the claw and the width—length ratio of the labium but it must be stressed that the
species under discussion belong to aberrant genera which are not typical of the subfamilies.
Acknowledgements
Most of the material discussed in this work is housed in the collections of the British Museum
(Natural History). Further specimens including holotypes and type-material have been kindly
made available for study by Mr Charles Chia-chu Tao of the Taiwan Agricultural Research
Institute, Taipeh; Dr J. P. Duffels of the Instituut voor Taxonomische Zodlogie (Zodlogisch
Museum), Amsterdam; Dr P. H. van Doesburg of the Rijksmuseum van Natuurlijke Historie,
Leiden; Dr Khoo Soo Ghee of the University of Malaya, Kuala Lumpur; Dr A. Tranfaglia of
the Istituto di Entomologia Agraria dell’Universita di Napoli, Portici and Dr D. R. Miller of the
USDA, Beltsville, who sent specimens from the United States National Museum, Washington.
The writer is much indebted for useful information on aphids given by Dr V. F. Eastop and
Dr R. E. Blackman of the British Museum (Natural History).
Abbreviations of the type-depositories mentioned in the text are as follows.
BMNH, London British Museum (Natural History), London
IEAUN, Portici Istituto di Entomologia Agraria dell’Universita di Napoli, Portici
ITZ, Amsterdam Instituut voor Taxonomische Zodélogie, Amsterdam
RNH, Leiden Rijksmuseum van Natuurlijke Historie, Leiden
TARI, Taipeh Taiwan Agricultural Research Institute, Taipeh
UM, Kuala Lumpur University of Malaya, Kuala Lumpur
USNM, Washington United States National Museum, Washington
Taxonomic treatment
An ideal classification of Coccoidea should be based on the taxonomy of adult males but so few
males have been studied that identification is possible only with adult females which are neotenic.
Female Pseudococcidae usually have three immature instars and males two immature and two
pupal instars. There is often little to distinguish immatures from adult females in life apart from
size. It is the experience of the writer, who has been engaged in identification of Coccoidea for
many years, that identification of immature stages is often just as important to the collector as
identification of the adult. When a new species is named, the name applies to all stages, not just
to the stage mentioned in the description. In the present work available immature stages are
discussed along with adult females.
13
PSEUDOCOCCINAE
ALLOMYRMOCOCCINI trib. n.
Type-genus: Allomyrmococcus Takahashi, 1941.
Pseudococcinae with body of adult female variously shaped, turbinate, or with abdomen rounded or
abruptly narrowed at third segment. Head either membranous or heavily sclerotized when there is a deep
constriction at about the prothorax. Anal lobes either wanting, in which case the anal lobe setae are short,
or well developed with long anal lobe setae about as long as body. Anal ring usually sclerotized, terminal
or situated a short distance from apex, with 6 setae but without pores except in one genus with minute
pore-like structures. Antennae 6-segmented, stout, well developed and often as long as body, of a distinc-
tive shape. The first segment on dorsum when mounted on slide, often large and wide; second segment
short and not articulating with third segment. Remaining segments long, widest at distal end except the
last segment, which is often depressed near middle of segment and variously curved. Fourth to sixth
segments with abundant short setae, the distance between setal bases shorter than length of a seta. First
and second segments with or without abundant setae. Longer and stouter sensory setae present on last
segment. Legs well developed, long, the tarsus much shorter than tibia, often with numerous setae.
Claw stout, each with a pair of flattened and expanded digitules, these about twice as long as claw and
about same width. Labium conical, with 3 segments and with basic number of 14 pairs of setae but some-
times with as many as 32 pairs. Ostioles situated on edges of body when mounted on slide or lying on
ventral surface, with an unusual development with heavily sclerotized and wide projecting lips, without
setae or with posterior lip in form of sclerotized flap with numerous setae, the inner edges of lips joined
by a membrane. Circulus present or absent. Body setae short, often abundant when they may cover the
entire body surface except for intersegmental areas. When less abundant they may be accompanied with
short blunt setae. Body pores often restricted to trilocular pores and simple pores but occasionally a few
oral collar ducts may be present in mid-thoracic and anterior abdominal segments.
DIscussION. Based on the characters of the adult female, the tribe is one of the most distinctive
in the family Pseudococcidae. As mentioned earlier in the section on classification, no adult
males have been studied to give a clue to the relationship of the tribe. The evidence suggests that
males exist and there is a pressing need for further observations in the areas where the tribe is
known.
DISTRIBUTION. Thus far the tribe occurs in Thailand, West Malaysia (Malaya), Sarawak,
Indonesia and Vietnam but the distribution may be extended after further study is made of nests
of the ants Hypoclinea and Polyrhachis.
Key to genera of Allomyrmococcini (adult females)
1 Body without recognizable anal lobes, posterior end of body rounded, narrow, with short
setae in normal positions of lobes. Body setae abundant on dorsal and ventral surfaces
PARAMYRMOCOCCUS (p. 54)
— Body with anal lobes, either poorly or well developed, with anal lobe setae about as long as body 2
2 Head heavily sclerotized and body constricted at prothorax. Long anal lobe setae and marginal
setae on at least first preceding segment. Ostioles with posterior lip in form of a sclerotized
flap which opens and closes against an apparently fixed sclerotized anterior lip. Anal ring
narrow with minute pore-like structures. ; . | MALAICOCCUS (p. 30)
- Head membranous, body not constricted at prothorax. Long setae on anal lobes only or, if
long marginal setae are present they are not on first preceding segment. Ostioles each with
two sclerotized lips, the posterior lip similar to anterior lip. Anal ring with broad sclerotized
rim without pores. 3
3 Anal ring situated a distance about its own length from apex of abdomen, ina membranous
depression in middle of sclerotized anal lobes. Body setae all pointed, densely covering
dorsal and ventral surfaces, the distance between setal bases shorter than length of one seta.
Legs about as long as body with abundant setae . ; . ALLOMYRMOCOCCUS (p. 15)
— Anal ring terminal, at apex of dorsal lobe-like structure between ventral anal lobes. Body setae
of two main types, one short and pointed and the other short and blunt, slightly clavate, the
distance between setal bases greater than length of single seta. Legs shorter than body length
with numerous setae but not densely covered; sometimes these setae are long and fleshy
HIPPEOCOCCUS (p. 20)
14
Key to genera of Allomyrmococcini (immature instars)
1 Body elongate-turbinate, without recognizable anal lobes, posterior end of body rounded with
short setae in normal positions of lobes. Body setae mainly short and cylindrical on dorsum
but with longer pointed setae on venter and around margins . PARAMYRMOCOCCUS (p. 54)
— Anal lobes developed to some extent even though poorly developed in some stages. Long anal
lobe setae as long as body . ; : 2
2 Head with some sclerotization even if only around the eyes, this sclerotization becoming more
extensive in each successive stage. Long marginal setae on at least first preceding segment
from anal lobes often as far forward as sixth preceding segment.
Body setae pointed but sometimes there are present also minute setae with the seta shorter
than diameter of setal base. Anterior ostioles with posterior lip in form of flap
MALAICOCCUS (p. 30)
-— Head always membranous. If long marginal setae are present they are not on first preceding
segment from anal lobes. : : 3
3 Anal ring situated at least its own length from apex of body i ina small membranous depression
in middle of sclerotized anal lobes. Legs about as long as body (first instar not seen)
ALLOMYRMOCOCCUS (p. 15)
- Anal ring terminal at apex of sclerotized dorsal lobe-like structure between sclerotized ventral
lobes. Legs long but shorter than body. Body setae pointed or long, fleshy and clavate
HIPPEOCOCCUS (p. 20)
ALLOMYRMOCOCCUS Takahashi, 1941
Allomyrmococcus Takahashi, 1941: 201. Type-species: Allomyrmococcus acariformis Takahashi, by
original designation and monotypy.
Only a single species is known so far. The genus has some peculiar characters among which are
the unusually long stout legs about as long as the body. The turbinate shape, the well-developed
anal lobes and dense covering of minute setae are some of the other most important characters.
There is little information on its biology except that colonies are covered with a dense cluster of
the ant Dolichoderus [=Hypoclinea] and the insects feed on young shoots. There is every indi-
cation that a honeydew channel or basket is formed between the anal lobes. The anal ring lies
some distance from the apex of the abdomen and the space between the lobes is in the form of a
concave depression which presumably retains honeydew.
DESCRIPTION. A broadly oval to turbinate body form with well-developed anal lobes each terminating
with a group of stout setae about as long as body. Anal ring sclerotized, without pores but with 6 setae
lying just outside the ring. Antennae 6-segmented, about as long as body, the second segment short and
not articulated with the third; all segments densely covered with slender setae. Legs stout, as long as
body or longer, covered with slender setae, tibia about four times as long as tarsus. Claw with a pair of
flat expanded digitules, these about as wide as claw at base and twice as long. Body densely covered with
minute pointed setae. Ostioles situated laterally, prominent, with well-developed sclerotized lips, without
setae or trilocular pores. Labium stout but longer than wide with 30 or more pairs of setae in adult
female, ventral or posterior surface with 2 pairs of posterior setae. Eyes present. Trilocular pores present
but sparse.
Immature instars similar to adult female in form but smaller and with fewer body setae. No first instar
available but second and third instars described here and illustrated.
DISTRIBUTION. At present the genus is known only from Thailand.
Allomyrmococcus acariformis Takahashi, 1941
Allomyrmococcus acariformis Takahashi, 1941: 201. Holotype 2, THAILAND: Mt Sutep, near Chiengrai,
on plant of family Leguminosae, associated with ant Dolichoderus sp. [= Hyploclinea sp.] (TARI,
Taipeh).
Allomyrmococcus acariformis Takahashi; Takahashi, 1942: 15.
Adult Female (Fig. 3)
DESCRIPTION. External appearance described originally as ‘Dark reddish brown and blackish with no
wax.’
15
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Fig. 3. Allomyrmococcus acariformis Takahashi. Adult female. Specimen from Thailand,
Mt Sutep.
16
When mounted on the slide a broadly oval to turbinate species about 1-85 mm long and 1-4 mm wide,
broad at anterior end and tapering posteriorly to a well-developed and protruding pair of rounded anal
lobes. Antennae 1-81—1-92 mm long thus about as long as body; with 6 segments which on the longest
antenna the first segment is 170 um long, the second 150 um, third 450 um, fourth 320 um, fifth 380 um
and sixth 450 um, the last segment slightly curved, narrow at base but becoming 65 um at widest point
a quarter of length from base before tapering to apex. Each segment densely covered with slender setae
which on first segment are about 32 um long and commonly 70 um long on last segment. These setae
also accompanied with a few long stout setae on apices of fifth and sixth segments. Legs quite long and
well developed. Hind trochanter + femur 800-810 zm long, hind tibia 750-850 zm long and hind tarsus
170-190 um long, often curved inwards at an angle to tibia. All segments including coxae densely covered
with slender setae, those on femur about 20 um long and on tibia about 48 um long. Claw stout 44 um
long, each with a pair of stout flat digitules about twice as long as claw. Labium about 200 um long but
width difficult to determine in available specimens. With 30-32 pairs of setae. Apical segment with
2 pairs ventral posterior setae, 1 pair minute apical setae, 4 pairs of subapical setae each 28 um long,
10 pairs of lateral and anterior setae the longest 40 um, 10-12 pairs of medial setae the longest 48 um,
3 pairs of basal setae the longest pair each 40 um long and 2 pairs each 24-32 um long. Clypeolabral
plate with about 80 setae. Anal ring on dorsal surface about twice its diameter from posterior end of body,
without pores but with 6 setae each about 80 um long. Ostioles large with thick heavily sclerotized and
protruding lips, without setae. Circulus absent. Anal lobes sclerotized on both surfaces, this sclerotization
extending forwards almost to level of vulva. Median dorsal surface between lobes membranous, depressed
and forming a ‘honeydew basket’. Each lobe with 6 dorsal and 4 ventral stout apical setae each about
1-5 mm long.
Body surface densely covered with slender pointed setae each about 16-20 um long, covering almost
entire surface except for intersegmental areas. Spaces between setal bases much shorter than length of one
seta, the spaces closer on dorsal surface than on ventral surface. The setae lie in various directions giving
the surface a wavy appearance. Setal bases of two sizes, the larger size much fewer. A pair of stout setae
each about 500 um long present on lateral edges of sixth abdominal segment. Trilocular pores sparse.
Third Instar Female (Fig. 4)
Similar to adult female but differing mainly in size. Body broadly oval and not so turbinate as in adult,
1-5 mm long and 1-25 mm wide. Anal lobes moderately developed. Antennae 6-segmented 1:35-1:42 mm
long, the longest antenna with first segment 150 um long, second 120 um, third 300 um, fourth 220 um,
fifth 270 um and sixth 360 um. All segments densely covered with slender setae but not so numerous as
in adult, the setae ranging in length from 24 um on first segment to 60 um on sixth segment. Legs as in
adult but shorter. Hind trochanter + femur 600-630 um long, hind tibia 550-570 um long and hind tarsus
150 um long. Claw stout, 40 um long with a pair of wide flat digitules about twice as long as claw. Labium
176 um long and 108 um wide, with 30-32 pairs of setae, distributed as in adult but differing in size.
Lateral setae each about 40 um long, longest anterior seta 32 um long, longest medial seta 40 um long,
3 pairs of basal setae, the longest pair each 40 um long and the other 2 pairs each 20 um long. Clypeo-
labral plate with about 55 setae. Ostioles well developed, as in adult. Anal ring on dorsum about twice
diameter from apex of body about 44 um wide with 6 setae each about 60 um long. Anal lobes sclerotized
on dorsal and ventral surfaces, each lobe with about 9 stout apical and subapical setae 1:55 mm long.
Median dorsal area between lobes forming a channel or ‘honeydew basket’. Body setae slender, abundant
and densely covering almost entire surface except for clear intersegmental areas, about 16-20 um long,
the spaces between setal bases much shorter than length of one seta, dorsal setae tending to be closer
together than ventral setae. A pair of lateral marginal setae present on sixth segment each about 400 um
long. Trilocular pores not numerous.
Second Instar Female (Fig. 5)
This instar has similar characters to those of adult and third instar female but can easily be distinguished
in possessing a pair of lateral setae on sixth abdominal segment each about 360 um long, a pair on dorsal
lateral edge of fourth segment each about 550 um long and a submarginal pair on dorsum of prothorax
each about 500 um long, the last 2 pairs with sclerotized areas around setal bases. Body ovoid, about
1-35 mm long and 1:0 mm wide, anal lobes moderately developed. Antennae 0-9 mm long, the first
segment 110 um long, the second 80 um long, third 180 um, fourth 130 um, fifth 160m and sixth
240 um long. Setae ranging from 24 um long on first segment to 36 um long on last segment. Legs with
hind trochanter+femur 450 um long, hind tibia 390 um and hind tarsus 150 um; all segments densely
covered with slender setae, those on disc of femur 16-20 um long and on disc of tarsus 32 um long. Claw
32 um long. Anal ring 40 um in diameter with 6 setae each 52 um long; situated about twice diameter
17
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Fig. 4. Allomyrmococcus acariformis Takahashi. Third instar female.
18
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Fig. 5 Allomyrmococcus acariformis Takahashi. Second instar female.
19
from apex of body. Labium 132 um long and 116 um wide, with numerous setae as in third and adult
instars but obscured in specimen available. Clypeolabral shield with numerous setae but difficult to
count in available specimen. Ostioles with well-developed sclerotized lips, the anterior pair lying on ventral
surface when mounted on slide. Anal lobes sclerotized, each with 11 apical and subapical setae about
1-5 mm long, their positions varying on dorsal and ventral surfaces but probably with 4 on dorsal surface
and 7 on ventral surface. Body setae slender, about 16 um long, covering almost entire surface except for
intersegmental spaces, the distance between setal bases shorter than length of one seta. Trilocular pores
few, scattered.
MATERIAL EXAMINED. THAILAND: 2 9, ‘Siam’, Mt Sutep, 5.iv.1940 (R. Takahashi; 1 third instar 2
and | second instar 2, same data as holotype (TARI, Taipeh).
This material has the same data as the holotype and it is assumed it is part of the original. The
specimens have been kindly made available for study by Mr Charles Chia-chu Tao of the Taiwan
Agricultural Research Institute, Taipeh.
HIPPEOCOCCUS Reyne, 1954
Hippeococcus Reyne, 1954 : 237. Type-species: Hippeococcus rappardi Reyne, by original designation.
This genus is known only from Java. It has some peculiar and unusual features connected with
its mode of life. According to Reyne the species are invariably associated with ants of the genus
Dolichoderus (=Hypoclinea). They are known to crawl on to ants’ backs and are carried away
when the ants are disturbed but large individuals may be carried in the ants’ jaws. Found on the
suckers and stems of the host, colonies are often covered with dense crowds of ants.
Reyne has given detailed descriptions of the species but there has been a pressing need for
modern illustrations. The following descriptions, therefore, serve to supplement Reyne’s and to
accentuate the principal characters.
In common with most mealybugs there are four female instars and all of these are described
for H. wegneri Reyne. Although Reyne mentioned only three instars it is possible that his descrip-
tions of the second instar may include characters of the third also. Only three female instars are
available here for H. rappardi Reyne; nevertheless, one or two specimens are at hand of what is
presumed to be the second instar male and this is described. Reyne described the adult female
before preparation as ‘Body gleaming, without a coating of wax’.
DESCRIPTION OF ADULT FEMALE. Body broadly turbinate, seventh and posterior segments narrow. Venter
of last segment folded on either side near the mid-line giving it the appearance, when viewed through the
stereomicroscope, of forming two parallel tubes, the apices forming the anal lobes containing 5—7 long setae
which may be almost as long as the body. The two tube-like parts of the last segment are covered dorsally
with a triangular lobe containing setae of various lengths and which bears the anal ring at the apex.
This ring is sclerotized, without pores, crescentic and contains 6 setae which increase in length and
thickness posteriorly. The anal ring is usually on level with apices of the anal lobes but in one species the
anal lobes are produced latero-posteriorly for a distance about same length again as last segment. Anten-
nae 6-segmented, first segment normal, second quite small and usually wider than long, both segments
with few setae. Third to sixth segments stout and elongate, containing numerous slender pointed setae,
the total length of antenna about half length of body. Ostioles well developed, lying at sides of body,
each with heavily sclerotized lips, the sclerotization surrounding anterior lips of first pair also extending
around eyes. Without membranous lips, setae or trilocular pores. Setae on both surfaces of body abundant,
mainly of two types, one of which is minute, clavate, usually 6-8 um long, almost transparent, and the
other type normal and pointed but short, about 20 um long. Other setae, present on the posterior abdo-
minal segments and legs, may be stout and long with a blunt tip or swollen apically and of a fleshy
appearance. Only the basal part of these setae is sclerotized, the remainder becoming more transparent
towards the apex. Claw stout, bearing a pair of quite wide and flat digitules each usually about as wide as
the claw and about twice as long. Trilocular pores present. Circulus oval.
DESCRIPTION OF IMMATURE INSTARS. Normally turbinate but becoming wider at each instar. Anal lobes
and dorsal triangular lobe containing anal ring present in all stages. Minute transparent clavate setae
absent. Short pointed setae absent in first instar but becoming more numerous in each successive stage.
Antennae of first instar with thickened long setae only. Second instar with abundant short setae on third
to sixth antennal segments and these setae become more numerous in successive instars.
20
Ls
from Java, Mt Kawa
imen
Adult female. Spec
Reyne
rdi
Hippeococcus rappa
Fig. 6
21
Key to species of Hippeococcus (adult females)
1 Anallobes projecting about the length of last segment from level of anal eae montanus Reyne (p. 22)
— Anal lobes terminating at about same level as anal ring F : 2
2 Ventral surface of abdomen with numerous stout or expanded setae in addition to long setae
on seventh and posterior segments. Anal lobes each with 6 long apical setae wegneri Reyne (p. 26)
— Ventral surface of abdomen without numerous expanded setae, short pointed setae present
only except for long setae on seventh and posterior segments. Anal lobes each with 5 long
apical setae : : : : : ; ; : ; : rappardi Reyne (p. 22)
Immature instars
A few instars have not been available for study and it is not possible to give a workable key.
The first instars of H. rappardi and H. wegneri have thick, fleshy and clavate setae on the dorsal
and ventral surfaces but in H. rappardi there are 2 long pointed setae on each side of sixth and
seventh segments. In H. wegneri there is 1 long seta and 1 short blunt seta in these positions.
The thick clavate setae persist in the second and third instars of H. wegneri but in H. rappardi
they are replaced by pointed setae. Instars are mainly distinguished by an increase in antennal
and leg lengths. The second instar males are easily recognized by the presence of multilocular
disc pores and oral collar tubular ducts.
Hippeococcus montanus Reyne, 1954
Hippeococcus montanus Reyne, 1954: 250. Holotype 2, JAvA [lost] (named from drawings in ITZ, Am-
sterdam).
There are no specimens available of this species as Reyne (1954) named it from certain drawings
made by P. van der Goot possibly in 1915. The illustration has the appearance of being accurately
executed. Although obviously a Hippeococcus species, it differs from H. rappardi and H. wegneri
in having quite long and prominent anal lobes that are set rather wide apart and protrude about
the same distance as the length of base or dorsal lobe covering them. In the other two species the
anal lobes are about the same length as the dorsal lobe.
Other than living in Java there is no further information on its locality but from the illustration
reproduced by Reyne, it should be easy to identify the insect when collected again.
Hippeococcus rappardi Reyne, 1954
Hippeococcus rappardi Reyne, 1954: 239. Holotype 2, JAVA: East, Yang Mts, Gondang Plantation, on
Ficus variegata, 21.vii.1950 (RNH, Leiden) [examined].
Adult Female (Fig. 6)
Body broadly turbinate, about 2:1 mm long and 1-45 mm wide, posterior segments tapering. Anal lobes
heavily sclerotized ventrally and with the dorsal surface forming lobe covering median area of last
segment, with anal ring at apex. Seventh and eighth segments sclerotized mid-dorsally, and areas of
sclerotization present around the well-developed ostioles and around the eyes. Entire surface with minute
sclerotized dots that give the body a slightly dark appearance even when not stained. Antennae about
1-55 mm long, the third to sixth segments densely covered with slender setae, those on the third segment
52-60 um long and those on sixth segment thicker and usually 100 um long. Legs well developed, hind
trochanter + femur 530 um long, hind tibia+ tarsus 540-560 um long, hind coxa with a few translucent
pores. Setae on legs of various sizes, on the hind tarsus they range from stiff pointed setae 40-100 um
long on the inner edge to stout fleshy setae, pointed, 52-160 um long on the outer edge. Claw about
44 um long. Labium 180 um long and 140 um wide with 14 pairs of setae. Apical segment with two pairs
of posterior setae, 1 pair of minute apical setae, 4 pairs of subapical setae each 32 um long, | pair lateral
setae each 52 um long and 2 pairs of anterior setae each 68 um long. Medial segment with a single pair
of medial setae each 60 um long and basal segment with 3 pairs basal setae, 1 pair 32 um long and 2 pairs
each 20 um long. Circulus oval, about 74 um wide. Anal ring at apex of abdomen, about 88 um wide,
with 6 setae, these becoming distinctly longer in a posterior direction, the posteriormost quite thick.
Dorsal setae of two main types. On all segments anterior to eighth there are numerous minute clavate
setae with a fleshy appearance, each about 6 um long. Other dorsal setae pointed, numerous and usually
26 um long. On the posterior abdominal segments the setae become longer. Trilocular pores not numerous.
22
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Fig. 7 Hippeococcus rappardi Reyne. Third instar female.
23
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Fig. 8 Hippeococcus rappardi Reyne. (A) First instar. (B) Second instar male.
24
Ventral surface with numerous minute blunt setae as on dorsum but they are absent between antennae
and on eighth and posterior segments. Other setae pointed, about 26 um long except for longer setae
near abdominal margins and for long setae 180-200 um long on the seventh and eighth segments. Tri-
locular pores tending to be more numerous than on dorsum but absent on head.
Third Instar Female (Fig. 7)
Body about 1-75 mm long and 0-75 mm wide, elongate, widest on thorax, abdomen tapering. Seventh
and eighth segments with distinct sclerotization in mid-regions. Anal lobes and dorsal median lobe
heavily sclerotized. Antennae 1-2 mm long, third to sixth segments with numerous short setae. Hind
trochanter+femur 450 um long, hind tibia+tarsus 460 um long. Claw about 36um long. Labium
148 um long and 100 um wide with same number of setae as in adult and about same size. Circulus oval,
56 um wide. Ostioles present at edges of body, similar to those of adult but smaller. Body setae short and
pointed only, with a similar distribution to those of adult female, but less numerous; mid-dorsal setae
about 40 um long and a typical short ventral seta about 40 um long. Seventh and eighth segments sclero-
tized over most of the area. On the seventh segment most of the setae lie in the sclerotized area and on the
eighth segment all the setae lie within the sclerotized area. Some long ventral setae on segments 7 and 8
stout and longer than the segments.
Second Instar Male (Fig. 8B)
The writer agrees with Reyne (1954) that this is the second instar male rather than the female because the
body has a vestiture of numerous tubular ducts and multilocular disc pores. Although second instar
females of other mealybugs may possess similar pores and ducts they usually become more numerous
in the third and adult instars. In this species they are absent completely in the third and adult female
instars. Numerous pores and ducts in the second instar male are present to secrete the pupal covering.
Body elongate, abdomen tapering, about 1-35 mm long and 0:55 mm wide. Antennae 800 um long, the
third to sixth segments densely covered with short slender setae. Legs well developed, hind trochanter +
femur 300 um long, hind tibia + tarsus 340 um long. Claw 28 um long. Labium 120 ym long and 100 pm
wide with 14 pairs of setae, common to the species. Dorsal labial setae shorter than those of previous
stages, subapical setae each 24 um long, lateral setae 36 um long, anterior setae each 40 um long. Medial
segment with medial setae 52 um long and basal segment with 1 pair basal setae 32 um long and 2 pairs
each 16 um long. Circulus oval, about 20 um wide. Ostioles well developed, on edges of body. Anal ring
64 um wide.
Dorsal setae all pointed, numerous, a mid-dorsal seta usually about 24 um long. Longer setae present
also on the lobe-like structure above anal lobes. Multilocular disc pores present, mainly in submedian
areas of thorax and anterior abdominal segments. Oral collar tubular ducts, each with the internal collar
flange-shaped and occupying about half length of duct, situated among the multilocular disc pores and
extending to margin. Trilocular pores sparse.
Ventral surface with numerous pointed setae tending to be longer than those on dorsum, a common
length being about 32 um. Two pairs of sclerotized areas on seventh and eighth segments each with a pair
of long pointed setae. Multilocular disc pores in mid-regions of thorax, anterior abdominal segments
and near spiracles. Tubular ducts present near the multiloculars and towards the margins. Trilocular pores
few.
First Instar (Fig. 8A)
Body narrowly turbinate, about 1-2 mm long and 0-4 mm wide at the thorax. Anal lobes and dorsal surface
forming lobe, heavily sclerotized. Anal ring about 60 um wide, without pores but with 6 setae at anterior
or dorsal half of the ring. These setae become progressively thicker and longer posteriorly, the longest
about 160 um. Anal lobes with 2 long dorsal setae and 3 ventral setae, each about 0-95 mm long. Antennae
6-segmented, about 630 um long with a few long setae slightly expanded distally, on each segment except
last where they are pointed. Legs well developed, hind trochanter + femur 220 um long, hind tibia + tarsus
about 310 um long. Claw 28 um long. Labium 120 um long and 88 um wide, the dorsal setae tending to
be shorter than in previous stage; subapical setae 24 um long, lateral setae 36 um long, anterior setae each
32 um long, medial setae 36 um long and basal setae with 1 pair 32 um long and 2 pairs each 12 um long.
Ostioles well developed, without setae but with lips heavily sclerotized. Circulus small and oval about
20 um wide.
Dorsal setae slightly expanded towards apex, each with a fleshy appearance and about 56 um long on
mid-abdomen where there are about 8 across a segment. Eighth segment with 2 pairs of long pointed
setae reaching beyond anal lobes. Trilocular pores few, forming a single submedian row and with one or
two around margins.
25
Ventral surface with setae similar to those on dorsum but tending to be longer, up to 60 um, except the
submarginal setae on seventh and eighth segments which are long and pointed, in pairs, on lightly sclero-
tized areas. Trilocular pores few, in submedian areas. Simple pores few on submargins, each about same
size as a trilocular pore but usually oval in shape.
MATERIAL EXAMINED. Hippeococcus rappardi Reyne, holotype 9, JAVA: East, Yang Mts, Gondang
Plantation (RNH, Leiden).
JAVA: Immatures with same data as holotype; immature instars, Mt Kawi, Gaden, on young
shoots of Litsea sp. (Lauraceae), ii.1951 (F. W. Rappard) (RNH, Leiden); 1 9 and immatures,
same data but labelled from Litsea sp., Eugenia sp. [Myrtaceae] and Rubus sp. [Rosaceae]
(BMNH, London).
Recorded also by Reyne (1954) from JAvA: Gondang, on Ficus variegata, on suckers of
Eugenia sp. and Litsea confusa; from Gaden, on stems of Rubus sp. and Eupatorium sp. (Com-
positae).
Hippeococcus wegneri Reyne, 1954
Hippeococcus wegneri Reyne, 1954: 255. Holotype 2, JAavA: West, Tjibodas, Botanical Garden, from
nest of Dolichoderus (= Hypoclinea) gibbifer Emery, i.1953 (RNH, Leiden) [examined].
Adult Female (Fig. 9)
Body broadly turbinate, attaining a length of 2:75 mm and a width of 2-0 mm, abdomen narrowing
abruptly. Anal lobes and dorsal surface of eighth segment heavily sclerotized. Sclerotization also in dorsal
median areas of fifth to seventh segments and in ventral submedian areas of seventh and eighth segments.
Anal lobe setae comprising three dorsal and four ventral, each about 1-5 mm long. Entire surface with
minute sclerotized dots giving the membranous body the appearance of being lightly sclerotized. Antennae
about 1-5 mm long with abundant slender setae on third to sixth segments. Setae on third segment about
60 um long, becoming progressively longer to sixth segment where they are thicker and 90-100 um long.
Legs well developed, hind trochanter+ femur 570-590 um long, hind tibia+tarsus 600-620 um long.
Claw 40 um long. Setae on legs of different shapes; they are long clavate and thick with a fleshy appear-
ance, or short or long and pointed, the longest on the outer side of tibia and tarsus. On the tarsus the
pointed setae are 40-160 um long and the stout fleshy setae 40-180 um long. Hind coxae with a few
translucent pores. Labium 184 um long and 120 um wide, with 14 pairs of setae. Apical segment with 2
pairs of posterior setae, a pair of minute apical setae, 4 pairs of subapical setae each 28 um long, a pair
of slender lateral setae each 48 um and 2 pairs of thick fleshy anterior setae each 80 um long. Medial
segment with 1 pair of fleshy setae each 92 um long and basal segment with 2 pairs of short pointed setae
each 20 um long and a pair of fleshy setae each 44 um long. Circulus about 68 um wide. Anal ring 84 um
wide, on level with apices of anal lobes, the anterior pair of setae 100 um long, the median pair 140 um
long and the posterior pair 180 um long, these setae becoming thicker as they become longer.
Dorsal surface anterior to seventh segment with numerous minute clavate setae each 6-8 um long, of a
fleshy appearance, barely taking up the stain. Short pointed setae each about 20 um long, evenly distri-
buted. Posterior abdominal segments with a few clavate or thick setae ranging in length from 30 to 120 um,
the dorsal lobe covering anal lobes with a few thick setae about 60 um long. Trilocular pores in moderate
numbers, evenly distributed.
Ventral surface with similar minute clavate and pointed setae as on dorsum, on sixth and anterior
segments. Long clavate or thick and bluntly pointed setae present across the abdominal segments ranging
in length from about 60 to 90 um except for some on seventh, eighth and anal lobe segments that may be
up to 260 um long. Trilocular pores numerous with an even distribution.
Third Instar Female (Fig. 10)
Body a similar turbinate shape to that of adult female but narrower, 2-0 mm long and 1-0 mm wide.
Sixth and posterior segments sclerotized with the 3 dorsal and 3 ventral anal lobe setae about 1-5 mm long.
Antennae 1-13 mm long, the third to sixth segments densely covered with short setae which become
longer towards the last segment. Hind legs with trochanter+ femur 450 um long, hind tibia + tarsus
480 um long. Claw about 30 um long. Labium 152 um long and 120 um wide, with 14 pairs of setae but
these shorter than in adult. Subapical setae each 26 um long, lateral setae each 36 um long, the 2 pairs
of anterior setae each 56 um long, thick and fleshy. Medial segment with medial setae thick, each 68 um
long and basal segment with 2 pairs of short pointed setae each 16 um long and 1 pair of thick setae
each 56 um long. Circulus oval, 68 um wide. Anal ring about 68 um wide. Body setae of two main types,
26
DJ.W.
Fig. 9 Hippeococcus wegneri Reyne. Adult female. Specimen from Java, Tjibodas.
27
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Fig. 10 Hippeococcus wegneri Reyne. Third instar female.
28
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VSIA fy 1)
fs / rail B
i / [i
A\VAveal
/P 7 Jj ifs \ \i j
ie a NT
j haf
Fig. 11 Hippeococcus wegneri Reyne. (A) First instar. (B) Second instar female.
5)
a clavate or expanded type, thick and fleshy, ranging in length from 50 to 100 um except at posterior
end of body where some are longer on ventral surface, and short pointed setae, which are numerous and
interspersed among the fleshy setae. The minute clavate setae are absent in this instar. Trilocular pores
sparse.
Second Instar Female (Fig. 11B)
Body narrowly turbinate, abdomen tapering, 1-4 mm long and 0-7 mm wide. Sclerotization at posterior
end of body on anal lobes, on dorsal surface of eighth segment and in median areas of seventh and sixth
segments. Legs with hind trochanter + femur 370 um long, hind tibia+ tarsus 390 um long. Claw about
28 um long. Antennae 900 um long, third to sixth segments with numerous slender setae. Ostioles less
developed than in previous stage. Circulus about 40 um wide. Labium with 14 pairs of setae. Apical
segment with 4 pairs of subapical setae each 24 um long, a pair of slender lateral setae each 36 um long
and 2 pairs of thick fleshy anterior setae each 48 um long. Medial segment with 1 pair of fleshy setae each
56 um long and basal segment with 2 pairs of slender setae 12 um long and a pair of thick setae each
44 um long. Body setae with similar distribution of clavate type to those in third stage but short pointed
setae less numerous especially on head and ventral thorax, the short dorsal setae about 24 um long and
the ventral setae about 32-40 um long. The clavate setae range in length from 50 to 100 um long except
at posterior end of body where some are 160 um long. Anal lobe setae difficult to measure but all are over
1-0 mm long. Trilocular pores sparse on both surfaces.
First Instar (Fig. 11A)
Body narrowly turbinate, about 1:35 mm long and 0-7 mm wide, anal lobes each with 3 dorsal and 2
ventral setae at apices, about 1-3 mm long. Anal lobes and dorsal lobe-like structure covering anal lobes
heavily sclerotized. Antennae 630 um long with clavate setae on all segments except last, which has long
pointed setae. Legs well developed, hind trochanter+femur 300 um long, hind tibia+tarsus 320 um
long. Claw 24 um long. Labium 108 um long and 80 um wide, with 14 pairs of setae, all slender except
for medial setae which are thick and fleshy. Apical setae each 24 um long, lateral setae each 28 um long,
anterior setae each 40 um long. Medial segment with medial setae 56 um long and basal segment with 2
pairs of basal setae 12 um long and 1 pair 32 um long. Ostioles small but with lips heavily sclerotized.
Circulus oval, about 20 um wide. Anal ring 44 um wide with 6 setae which become progressively longer
posteriorly, the second pair twice as long as anterior pair and posterior pair half as long again as median
pair.
Dorsal surface without short pointed setae but with long*clavate setae ranging in length from 40 to
68 um. Eighth segment with 4 stout pointed setae as long as anal lobes. Trilocular pores sparse.
Ventral surface with similar clavate setae to those on dorsum but some are longer, approaching 80 um
long. Long stout clavate to pointed setae present also on small sclerotized areas on seventh and eighth
segments. Trilocular pores few.
MATERIAL EXAMINED. Hippeococcus wegneri Reyne, holotype 9, JAVA: West, Tjibodas, Botanical
Garden, 1.1953 (A. M. R. Wegner) (RNH, Leiden).
JAVA: immatures, same data as holotype; paratypes and immatures, same data, 1 2 on young
shoots and fruit of Diospyros kaki [Ebenaceae] (RNH, Leiden), 1 2 (ITZ, Amsterdam); 6 9 and
immatures, i.1953 (A. M. R. Wegner); 2 9, vi.1953 (A. M. R. Wegner); 1 2 Tjibodas, 22.x.1973
(B. Bolton) (BMNH, London).
Recorded also by Reyne (1954) on Cyclanthera explodens (Cucurbitaceae).
MALAICOCCUS Takahashi, 1950
Malaicoccus Takahashi, 1950: 65. Type-species: Malaicoccus riouwensis Takahashi, by original desig-
nation.
This is an unusual genus of the Pseudococcidae in that the known species have the prothorax
constricted and most of the head is heavily sclerotized. In life they have a strong resemblance to
aphids and may often be mistaken for them. All are associated with ants of the genera Polyrhachis
and Hypoclinea but little is known of their life history or habits. The mealybugs are attended by
the ants which often carry them in their mandibles at the constriction on the prothorax.
DESCRIPTION OF ADULT FEMALE. Body form ovoid to globular with a definite constriction on prothorax
opposite first coxae. Head heavily sclerotized around antennal bases, the sclerotization often extending to
30
first pair of ostioles; the front of head on ventral surface almost completely sclerotized forward from
clypeus. Antennae 6-segmented, often longer than entire body and densely covered with slender setae;
the first segment arising from dorsal surface. Second segment short and not articulating with third seg-
ment. Legs well developed, often with numerous setae of different sizes. Claws stout, with flattened digi-
tules which are greater in size than claw itself. Labium conical with 21-32 pairs of setae, always with 2
pairs of posterior setae, a pair of minute apical setae, 4 pairs of subapical setae and 1 pair of lateral setae.
The supernumerary setae occur in the anterior and medial groups. Circulus present, usually well devel-
oped with deep lateral constrictions. Ostioles of a distinctive type, the anterior pair each with an apparently
fixed anterior lip which is arc-shaped. A heavily sclerotized semi-circular lower lip containing a few setae
is hinged to close over to the anterior lip. These ostioles often lie on ventral surface on slides due to con-
vexity of body or more often on margins so that they are viewed in lateral aspect. Posterior ostioles with
anterior lip apparently fixed and with the posterior lip narrower than in anterior ostiole, with or without
small setae. Anal ring terminal, narrow, with a few minute pores and 6 setae. Two crescentic or triangular
areas of sclerotization containing setae present at either side of anal ring or just anterior to it. The anal
ring is depressed behind the crescentic areas so that apex of abdomen forms a small ‘honeydew basket’.
Setae on margins of last 2-4 segments about as long as body and often accompanied with 1 or 2 short
setae. Body setae minute, always numerous and sometimes densely covering the surface in which case the
distance between the setal bases is shorter than the length of a seta. Other setae stout, of various lengths.
Posterior end of body with sclerotized areas on both sides of anal lobes and on ventral surface at margin
of some preceding segments.
DESCRIPTION OF IMMATURE INSTARS. Body oval but head angular in first instar and becoming more rounded
in successive instars. Prothorax with a slight constriction in first instar but constriction increasingly
apparent in second and third instars. Sclerotization on head starting around eyes in first instar and
increasing in each instar to around base of antenna and in an area forward from clypeus. Anal lobes
always developed to some extent. Antenna 6-segmented, often as long as body. In the first instar long
pointed setae form whorls usually at distal end of segments but in the second and third instars the segments
become densely covered with short pointed setae. Legs well developed, claw stout, with 2 flat, wide
digitules even in the first instar. Labium with 14 basic pairs of setae but this number often increasing in
third instar. Ostioles similar in shape to those of adult but without setae. Anal ring usually narrow with
a few minute pores. Setae about as long as body on anal lobes and on at least first segment preceding
lobes, often present as far forward as second segment or sixth preceding Jobes, always on sclerotized areas.
Body setae long on first instar but second and third instars with increasing numbers of short setae. In
some species there are numerous minute setae with the seta shorter than diameter of setal base. Trilocular
pores and simple pores present only except in second instar male which has multilocular disc pores and
oral collar tubular ducts also.
Discussion. The genus is probably more closely related to Allomyrmococcus than to any other,
differing mainly in the sclerotized head, the presence of a circulus and the terminal anal ring.
DISTRIBUTION. Malaya and the Riouw Is in Indonesia. A third instar female of an undescribed
species is known among material of Allomyrmococcus acariformis collected by Takahashi in
Thailand. See also the record from Sarawak on p. 6.
Key to species of Malaicoccus (adult females)
1 A distinct tuft of short slender setae opposite each first spiracle, the space between these setal
bases shorter than the length of a seta . ; : : ; 2
— Tuft of short setae opposite each first spiracle absent 5 formicarii Takahashi (p. 33)
2 Setae in median area of ventral thorax all short and slender except for an occasional long seta
khooi sp. n. (p. 39)
— Setae in median area of ventral thorax with numerous long stout setae in addition to short
slender setae. ; ‘ 3
3 Long marginal setae each nearly 1: 0 mm long c on last 4 segments. Long setae in median areas
of anterior ventral abdominal segments absent. : moundi sp. n. (p. 44)
— Long marginal setae nearly 1-0 mm long on last 2 segments only. Marginal setae on preceding
segments much shorter. Long setae present in median areas of all abdominal segments. 4
4 Hind tibia with setae 40-60 um long, the spaces between setal bases often longer than a seta.
A seta 80 um long on seventh segment in addition to long setae on eighth segment and anal
lobes . , . : ; : : A : : j takahashii sp. n.(p. 52)
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Fig. 12 Malaicoccus formicarii Takahashi. Adult female. Specimen from Malaya, Fraser’s Hill.
32
— Hind tibia with numerous setae about 28 um long, the spaces between setal bases shorter than
a seta. Single setae 200-400 um long on margins of sixth and seventh segments in addition
to long setae on eighth segment and anal lobes. : : riouwensis Takahashi (p. 50)
Immature Instars
It is hoped that the immature instars described here may be recognized easily from the illustra-
tions and descriptions but in the absence of suitable material of some species it is not possible
to present a workable key. Some of the following distinguishing characters may prove useful.
In M. takahashii there are long setae on the anal lobes and margins of the eighth abdominal
segment only but in the other species studied, long marginal setae reach as far forward as the
second abdominal segment or sixth preceding segment from anal lobes. M. khooi, M. moundi and
M. takahashii have unusually short setae in the second and third instars, each seta much shorter
than the diameter of the setal bases. The characters of the adult female are reflected in the
immature instars. For instance, in M. khooi there is a short wide terminal segment on the
antenna in the adult, third and second instars and there is an absence of long setae on the ventral
surface of thorax in the adult and third instars. The labium of the first and second instars has the
basic number of 14 pairs of setae. In M. moundi this number persists in the third instar but in the
other species studied the number increases to as many as 18 pairs in the third instar. The labium
of the second and third instars of M. khooi has a primary medial seta which is clavate and digitule-
like. Second instar males are easy to recognize by the multilocular disc pores and oral collar
tubular ducts on both surfaces.
Malaicoccus formicarii Takahashi, 1951
Malaicoccus formicarii Takahashi, 1951: 5. LECTOTYPE 9, West MataysiA: Malaya, Fraser’s Hill,
6.vi.1943 (R. Takahashi) (BMNH, London), here designated [examined].
Adult Female (Fig. 12)
External appearance described as ‘brownish black, without wax’.
On the slide body quite rotund except for a constriction on prothorax. Head and thorax heavily
sclerotized around bases of antennae, in a broad band posterior to first pair of ostioles and ventrally in
an area forward from clypeus to margins, there being a separate area running from the dorsal to ventral
surface enclosing eye. The extent of the sclerotization varies slightly in different specimens. Antennae
2:3-2:5 mm long, the second segment longer than wide and all segments densely covered with slender
setae which become longer towards apical segment. In an antenna 2:45 mm long the first segment is
300 um long, second 140 um, third 320 um, fourth 340 um, fifth 370 um and sixth 950 um, the last
segment strongly curved. Legs well developed, hind trochanter + femur 610-670 um long, hind tibia +
tarsus 670-760 um long, always longer than trochanter+ femur. Hind coxae with translucent pores to-
wards outer edge. All segments densely covered with short setae. Claw 48 um long. Circulus 150 um
wide, wider than long with a constriction on either side. Anal ring elongate and becoming quite narrow
at the point of constriction, easily distorted, with a few minute pores. Ostioles well developed, the anterior
pair each with a narrow sclerotized lip which is apparently fixed, the posterior lip in the form of a hinged
plate containing short setae which, when not flat against anterior lip, opens to almost right-angles to body.
Posterior pair similar but posterior lip narrower, with a few setae. Labium 210 um long and 120 um
wide, with 23-30 pairs of setae. Apical segment with 2 pairs of posterior setae, a pair of minute apical
setae, 4 pairs of subapical setae each 28 um long, 1 pair of lateral setae each 72 um long, 3-7 pairs of
anterior setae, the longest 60 um. Medial segment with 9-12 pairs of setae, the longest 72 um long. Basal
segment with a pair of short setae each 24 um long and a pair each 32 um long. Body setae all pointed.
Long marginal setae in four distinct groups on either side of abdomen, all about 1-0 mm long. Anal lobe
setae comprising a pair on dorsal surface on a small triangular area of sclerotization and a ventral pair
on a larger subrectangular sclerotized area. A pair of long setae also present on ventral margin of eighth
segment at base of large sclerotized area. A pair also located on margin of seventh segment and a single
seta on margin of sixth segment. Dorsal setae abundant, covering most of surface except parts of thorax
and head where there are naked areas. The setae lie in definite waves or directions on each segment, a
common length being about 16 um except for longer setae on posterior abdominal segments and on two
sclerotized crescentic areas anterior-lateral to anal ring. Ventral setae also abundant but not so numerous
as on dorsum, mainly 20 um long. Trilocular pores very few on both surfaces except for a noticeable
concentration near inner edge of first coxa and around the labium. A few minute circular pores also present,
each with a diameter much smaller than a trilocular pore or setal base. Tubular ducts absent.
52
ih hey Ps / jie gaa
ei pit hiya eh
epee NOLIN
om aval [A 4
CL : i! va ! iy \
fai rR | | NS
\
|
.
f
D.J.W.
Fig. 13. Malaicoccus formicarii Takahashi. Third instar female.
34
eat x
; y ye Ys I) 8
Yi.’
a “il \
D.J.W.
Fig. 14 Malaicoccus formicarii Takahashi. Second instar female. Specimen from Selangor,
Ula Langat.
35
oy
D.J.W.
Fig. 15 Malaicoccus formicarii Takahashi. First instar.
36
Pp
> 2»
ok
ey vt w ys
WS y-
ae eae tae
ile!
,
a: AN ra
Ki it
Ne me y ( A
Ni if ti
Fig. 16 Malaicoccus khooi sp. n. Adult female. Specimen from Malaya, Genting Highlands,
on Poikilospermum cordifolium.
37
Third Instar Female (Fig. 13)
This instar has many of the features of the adult female but the body is less broadly oval, 1-75 mm long
and 1:2 mm wide and with a smaller constriction opposite first coxae. Antennae 1:70-1:75 mm long, the
first two segments with numerous short setae and the third to sixth segments densely covered with slender
setae. Long marginal setae on abdomen numbering 6 pairs, about 1-0 mm long on irregular sclerotized
areas. Short body setae less numerous than in previous stage, the dorsal setae 20-28 um long and some
of the ventral slightly longer. Other body setae present much longer than the dorsal, ranging from 60 to
150 um long and ventral setae distributed across the segments 60-180 um long. Sclerotization not so
extensive as in adult. In addition to areas around marginal setae and behind the antennae there is a
distinctive sclerotized area just below first pair of ostioles, and two triangular areas between antennal
bases on dorsum, these containing setae. Circulus about 88 um wide with a constriction on either side.
Trilocular pores and smaller simple pores present but few. Anal ring narrow with lateral constrictions and
becoming distorted during the preparation of the insect on the slide, with a few minute pores. Legs with
a few short setae and some long pointed setae, hind trochanter + femur 430-450 um long, hind tibia +
tarsus 500-510 um long. Claw 44 um long. Labium 160 um long and 100 um wide, with 17-18 pairs of
setae. Apical segment with 4 pairs of subapical setae each 28 um long, a pair of slender lateral setae each
36 um long and 2-3 pairs anterior setae, the longest 68 um. Medial segment with 3-4 pairs, the longest
68 um long, basal segment with 2 pairs basal setae each 24 um long and a pair each 60 um long.
Second Instar Female (Fig. 14)
Body oval, about 1-35 mm long and 0-8 mm wide, with anterior end angled and with slight constrictions
on the thorax opposite the first coxae. Antennae about 1-0 mm long, with long pointed setae and with a
few short slender setae on first segment and numerous such setae on third to sixth segments. Legs with
hind trochanter + femur 320 um long, hind tibia+tarsus 350 um long. Claw about 36 um long. Labium
130 um long and 80 um wide with 14 pairs of setae. Apical segment with subapical setae each 24 um long,
a pair of slender lateral setae each 32 um long and 2 pairs of anterior setae, the longest 72 um long.
Medial segment with a single pair of medial setae each 56 um long and basal segment with 3 pairs of
setae, a long pair each 52 um long and 2 shorter pairs each 20 um long. Anal ring sclerotized with 6 setae
about 80 um long, with a few minute pores at anterior end and one or two only at sides. Circulus 64 um
wide, oval with a slight notch on either side. Marginal setae on abdomen reaching over 1-0 mm in length.
On the anal lobes there are 2 dorsal and 2 ventral setae, the dorsal setae on an irregular-shaped area of
sclerotization and the ventral pair on a larger area that extends inwards over most of segment. The
anterior long marginal setae either on venter or dorsum when mounted on slide depending on convexity
of body. Each long seta accompanied with a much shorter seta, their bases on an irregular-shaped sclero-
tized area. Other sclerotized areas at bases of shorter setae on thorax, behind the antennae and on either
side of anterior end of anal ring, the latter areas crescentic or triangular containing a few setae. Other
body setae less numerous than in third stage, thick pointed setae present up to 120 um long, also short
pointed setae which on dorsum are 16-20 um long and on venter about 40 um long. Ostioles well devel-
oped, the anterior pair with a movable hinged lip.
First Instar (Fig. 15)
Body oval, about 1-05 mm long and 0-6 mm wide. Anal lobes poorly developed, anal ring on apex of
abdomen, projecting further than anal lobes. Anal lobes with 2 ventral apical setae about 1-1 mm long,
1 dorsal seta about same length and another about half the length. The five preceding segments each with
marginal ventral setae, these also about 1-0 mm long, all of these setae including those on anal lobes, on
irregular-shaped sclerotized areas which, anterior to anal lobes, also contain a much shorter seta. Sclero-
tized areas present on head around the eyes and in a crescentic patch on either side of anal ring. Antennae
620 um long, with long pointed setae only on each segment. Ostioles prominent with heavily sclerotized
lips. Circulus apparently absent. Anal ring sclerotized with 6 setae each about 44 um long and a few
minute pores at anterior edge only. Legs well developed, hind trochanter+femur 220 um long, hind
tibia + tarsus 260 um long. Labium 110 um long and 70 um wide with 14 pairs of setae. Subapical setae
each 20 um long, a single pair of lateral setae each 40 um long and 2 pairs of anterior setae the longest
60 um long. Medial setae present as a single pair each 64 um long. Basal segment with 2 pairs of setae
each 20 um long and a pair 44 um long. Trilocular pores and simple circular pores in submedian areas,
quite sparse.
MATERIAL EXAMINED. Malaicoccus formicarii Takahashi, lectotype 9, West MALAYSIA: Malaya,
Fraser’s Hill, 6.vi.1943 (R. Takahashi) (BMNH, London).
38
West Mataysia: 17 9 and immatures, same data as lectotype (BMNH, London); 7 9, Selangor,
Kepong, on Bauhinia sp. [Leguminosae], 29.viii.1943 (R. Takahashi); 2 2 and immatures, Selan-
gor, Ula Langat, on ‘jungle plant’, 1928 (H. 7. Pagden); immatures, Fraser’s Hill, on Languas sp.
[=Alpinia] [Zinziberaceae], 6.vi.1943 (R. Takahashi) (BMNH, London); 3 9, Cameron High-
lands, on Rubus sp. [Rosaceae], x.1944 (R. Takahashi) (UM, Kuala Lumpur); 2 2 and immatures,
Gombak, on Mikania scandens [Compositae], 4.xi.1971 (Y. P. Tho) (UM, Kuala Lumpur).
Discussion. The adult female is unlike any other in the genus in lacking a tuft of short setae on
the venter opposite the first spiracles and in having the short body setae so densely covered that
the distance between setal bases is shorter than the length of a seta.
Malaicoccus khooi sp. n.
Adult Female (Fig. 16)
External appearance not known. Body broadly oval, about 2-35 mm long and 2:17 mm wide with a deep
constriction opposite first coxae. Head heavily sclerotized around antennal bases on dorsum and en-
circling the eyes, this area continuing to ventral surface just above anterior ostioles. Ventral head region
anterior to clypeus almost completely sclerotized. Antennae about 1:88 mm long, first segment 250 um,
second 130 um, third 360 um, fourth 300 um, fifth 260 um and sixth 560 um long, the last segment rather
short and squat compared with that in other species. All segments densely covered with slender setae.
Labium 208 um long and 108 um wide with 27 pairs of pointed setae. Apical segment with a pair of
posterior setae, a pair of minute apical setae about 4 um long, 4 pairs of subapical setae each 32 um long,
a pair of lateral setae 52 pm long and 2 pairs of anterior setae, the longest 72 pm. Medial segment with
11 pairs of setae, the longest 64 um long. Basal segment with 3 pairs of setae, the basal pair 16 um long
and the longest 56 um. Circulus rather large with a deep constriction at either side, about 230 um at its
widest at posterior end. Legs stout, well developed, hind trochanter+ femur 520 um long, hind tibia +
tarsus 460 um long and thus shorter than trochanter + femur. Hind coxa with a few translucent pores on
outer dorsal surface, short setae on outer ventral surface and longer pointed setae on inner ventral
surface. Long setae also present on outer margin of trochanter and a few present on femur, tibia and
tarsus: a few short setae also present on proximal end of femur. Claw 40 um long. Anal ring reduced to
a narrow rim with 6 setae each about 64 um long, and a few minute pores. Ostioles well developed, the
anterior pair lying on ventral surface when mounted on the slide due to convexity of body. Both pairs
with edge of anterior lip heavily sclerotized and probably fixed, the posterior lips forming a hinged
semi-circular plate which is capable of folding over to anterior lip, this plate with numerous short setae
on anterior ostioles but without setae in posterior pair.
Marginal setae on abdomen long, on anal lobes and three preceding segments. Anal lobes with 1 long
dorsal and 2 long ventral setae each 850 um long, the dorsal surface also with a shorter seta about one-
third length on an elongate-oval area of sclerotization. Ventral anal lobe setae at apex of a subrectangular
sclerotized area. Marginal setae on three preceding segments slightly ventral in position. Eighth segment
with a single seta 950 um long and a much shorter seta on a subrectangular area of sclerotization. Seventh
segment with a seta 480-550 um long and a shorter seta on a small sclerotized patch. Sixth segment with
a seta 490-600 um long and also a short seta on a small sclerotized patch. Dorsal surface of head lateral
to each antennal base with a long pointed seta about 32 um long. A few other setae 12-18 um long also
within the sclerotized areas of head, on abdominal margins and in mid-regions of eighth segment on dor-
sum and seventh and eighth segments on venter. A few setae present on a sclerotized crescentic area on
either side of anal ring. Other body setae numerous, ranging from 8 to 16 um long on dorsum, the larger
setae easily noticeable by the larger setal bases, evenly distributed except on head and thorax where there
are a few completely bare areas. Ventral body setae usually longer than dorsal, 12-20 um long, the longer
setae being the most numerous. A distinct tuft of short setae present near margin opposite each first
spiracle. Trilocular pores not numerous, evenly distributed but a few concentrated near labium. Minute
circular pores sparse, each with a diameter less than a setal base. Tubular ducts each with a diameter
smaller than a trilocular pore, represented by one or two on inner side of second and third coxae.
Third Instar Female (Fig. 17)
Body similar in shape to that of adult female with deep constriction opposite first pair of coxae, 1-4 mm
long and 0-9 mm wide. Antennae 1:25-1:30 mm long, the longest with first segment 180 um long and
160 um wide, second 90 um long, third 260 um, fourth 230 um, fifth 200 um and sixth 430 um. All seg-
39
Fig. 17. Malaicoccus khooi sp. n. Third instar female.
40
Fig. 18 Malaicoccus khooi sp. n. Second instar female.
41
Fig. 19 Malaicoccus moundi sp. n. Adult female. Specimen from Malaya, Genting Highlands,
on stems of Uncaria sp.
42
ments densely covered with slender setae ranging in length from 30 to 40 um on first segment to 45 um
on last segment. Legs well developed, stout. Hind trochanter + femur 390-400 um long, hind tibia + tarsus
360-370 um long. All legs with pointed setae of different sizes but second and hind legs each with a pair
of knobbed digitule-like setae on outer proximal end of tibia, the first legs with similar setae at distal end
of femur and along outer edge of tibia. Labium 160 um long and 100 um wide, with 17 pairs of setae.
Apical segment with a pair of posterior setae, 1 pair of minute apical setae, 4 pairs of subapical setae
each 24 um long, | pair of lateral setae each 44 um long, 2 pairs of anterior setae the longest 68 um long.
Medial segment with 4 pairs of setae, one of the pairs knobbed and digitule-like, 48 um long. Basal
segment with 2 pairs 16 um long and 1 pair 52 um long. Circulus constricted, widest at posterior end,
120 um wide. Anal ring slender with a few minute pores and 6 setae each 64 um long. Ostioles as in adult
female, the anterior pair lying on ventral surface when mounted on slide but these ostioles without setae.
Anal lobes sclerotized on both surfaces. Dorsal surface of each lobe with a seta 0:85 mm long and a shorter
330 um long. Ventral surface of lobe with apical seta 0-85 mm long and another probably shorter but
broken in available specimens. Stout marginal setae on 6 anterior segments. On the eighth segment
these setae are 0-85 mm long, on the seventh 350 um, sixth 450 um, fifth 450 um, fourth 500 ym and third
550 um long.
Dorsal surface with setae of different sizes. A pair of stout marginal setae each 350 um long present
just behind first antennal segment. Other setae about 50-60 um long, few on prothorax, head and last
two abdominal segments and singly on margins of most segments. On last segments they are present on
two crescentic sclerotized areas anterior-lateral to anal ring. Short slender setae numerous, each 6-8 um
long. Minute setae abundant, each with seta much shorter than setal base. Ventral setae of similar sizes
to those on dorsum but a pair 300 um long on head margin. Other long setae, the longest of which are
85 um, present in transverse rows on abdomen, head and prothorax. Slender setae 4-12 um long, few on
abdomen but numerous in median areas of head and thorax. Minute setae present in moderate numbers,
each much shorter than setal base. Trilocular pores and simple circular pores not numerous, scattered.
Second Instar Female (Fig. 18)
Body oval, about 1-0 mm long and 0:6 mm wide, sclerotized on head around eyes and on a few setal
bases. Antennae 840-880 um long. On longest antenna first segment 110 um long, second 60 um, third
160 um, fourth 150 um, fifth 120 um and sixth 280 um. Slender setae numerous but on each of first and
second segments there are one or two knobbed digitule-like setae. Legs well developed with hind tro-
chanter + femur 270-280 um long, hind tibia+ tarsus 270 um long. Knobbed setae present at distal end
of first and second femora and on outer edge of all tibiae. Other leg setae all pointed except for tarsal
digitules. Claw stout 28 um long. Labium 120 um long and 80 um wide, with 14 pairs of setae. Apical
segment with minute pair of apical setae, 4 pairs of subapical setae each 20 um long, | pair of lateral
setae each 32 um long and 2 pairs of anterior setae, the longest 56 um long. Medial segment with 1 pair
of knobbed setae 52 um long. Basal segment with 2 pairs of short setae 12 um long and 1 pair 36 um
long. Circulus constricted, 68 um wide. Ostioles rather large for size of body, the anterior pair lying
behind eye on ventral surface when mounted on slide, without setae. Anal lobes sclerotized, each with a
dorsal seta 0-8 mm long, another 270 um long and a ventral apical seta 0-8 mm long with another 450 um
long. Anterior six segments with long marginal setae, the bases surrounded by sclerotized areas. Eighth
segment with marginal setae 0-8 mm long, seventh 250 um, sixth 350 um, fifth 490 um, fourth 550 um
and third 550 um long. Anal ring with a few minute pores and 6 setae each 60 um long.
Dorsal surface with stout setae on thorax and posterior abdominal segments 24-50 um long. On re-
mainder of abdomen the setae are small and slender, about 12 um long. A pair of long setae behind first
antennal segment each about 210 um long, the area about the setal bases sclerotized. Minute setae
numerous, each with seta much shorter than diameter of setal base. Ventral setae of different sizes, the
longest about 50 um accompanied with short slender setae and a few of the minute type. Trilocular pores
and simple circular pores sparse.
Holotype 9, West MALAysIA: Malaya, Genting Highlands, on Poikilospermum cordifolium
[Urticaceae], attended by ant Hypoclinea sp., 4.x.1973 (L. A. Mound) (BMNH, London).
Paratypes. WesT MALAysIA: 3 9 and immatures, same data as holotype; 1 2, Genting High-
lands, on low herbaceous plant, attended by ant Hypoclinea sp., 4.x.1973 (B. Bolton) (BMNH,
London); 2 9, Gombak F. S., on bamboo [Gramineae], 26.x.1971 (Khoo Soo Ghee) (UM, Kuala
Lumpur).
Discussion. This species is similar to M. moundi in having setae about 1-0 mm long on anal lobes
and margins of the three preceding segments but it differs in the setae on mid-ventral thorax
43
which are short and slender, rarely more than 20 um long whereas in M. moundi there are numer-
ous setae up to 130 um long in this area. The sixth antennal segment in M. khooi is about 560 um
long but in M. moundi it is more slender and 710 um long.
Malaicoccus moundi sp. n.
Adult Female (Fig. 19)
Body broadly oval, with a distinct constriction on margins opposite first coxae, about 1-75 mm long and
1-15 mm wide. Head sclerotized around antennal bases on dorsal surface and over entire apex of venter
as far as clypeus and almost to constriction on margin. Antennae densely covered with short setae which
become longer towards last segment. Total length of antennae 2:12 mm and longer than body in available
specimen; first segment 260 um long, second 120 um, third 370 um, fourth and fifth each 330 um and sixth
segment 710 um long. Legs well developed, hind trochanter+femur 640 um long, hind tibia+ tarsus
630 um long hence slightly shorter than trochanter+ femur. All segments with moderately long setae,
hind coxa with a few translucent pores on dorsal outer half. Claw 56 um long. Labium 212 um long and
124 um wide, with 22 pairs of pointed setae. Apical segment with 2 pairs of posterior setae, 1 pair of apical
setae about 16 um long, 4 pairs of subapical setae each about 28 um long, | pair of slender lateral setae
each 64 um long and 2 pairs of anterior setae, the longest 92 um. Medial segment with 8 pairs of setae,
the longest 84 um. Basal segment with 4 pairs of setae, the shortest 22 um long and the longest 68 um
long. Circulus large with a deep constriction on either side, the posterior edge wider than anterior edge,
240 um at widest. Anal ring elongate and narrow, constricted, but distorted in available specimen; with
6 setae, each 72 um long and a row of minute pores. Ostioles well developed, each with a fixed sclerotized
anterior lip and a hinged sclerotized posterior lip which may completely close the ostiole. Anterior pair
on ventral margin on slide probably due to convexity of body, the posterior lips with a few setae.
Marginal setae on abdomen, on anal lobes and three preceding segments, the long setae all about
1-2 mm long. Anal lobe setae represented by 2 on dorsum, one thinner than the other, at apex of an
elongate area of sclerotization. Ventrally each lobe with a pair of long setae and a large subrectangular
area of sclerotization. Sixth to eighth segments with long setae on ventral margins, those on sixth segment
accompanied with a shorter seta about 130 um long, on the seventh segment with a shorter seta about
130 um long and on the eighth segment with a seta about 400 um long. Stout setae 50-130 um long present
dorsally on head, prothorax, abdominal margins, median areas of eighth and posterior segments and in
a cluster on a sclerotized crescentic area on each side of anal ring. Similar setae present ventrally on head,
in median areas of thorax and in mid-region of seventh and posterior segments of abdomen. One or two
longer setae 130-300 um long, in median area of dorsal and ventral eighth abdominal segment. Body
setae abundant except on dorsum of head and prothorax where there are extensive bare areas. These
setae extremely slender, 6-24 um long, the majority being the smallest. Ventral surface opposite first
spiracles with a distinctive tuft of short setae. Trilocular pores not numerous, evenly distributed but
absent on head; there are, however, noticeable concentrations around labium. Simple circular pores,
each with a diameter smaller than a trilocular pore, in moderate numbers between second and third coxae.
Third Instar Female (Fig. 20)
Body broadly-oval with a constriction opposite first coxae giving the body a pyriform appearance, about
1-4 mm long and 0-95 mm wide. Head sclerotized on dorsum between antennal bases, on venter between
clypeus and anterior margins, and surrounding eyes. Anal lobes moderately developed. Antennae about
1:57-1:58 mm long, the first segment 170 um long, second 100 um, third and fourth each 250 um, fifth
260 um and sixth 550 um long, each segment with numerous slender setae which on first segment are
about 32 um long and on sixth segment about 36 um long. Legs with hind trochanter + femur 460-470 um
long, hind tibia + tarsus about 460 um long, stout. Labium 160 um long and 100 um wide. Apical segment
with 2 pairs of posterior setae, a pair of minute apical setae, 4 pairs of subapical setae each 24 um long,
1 pair of lateral setae each 40 um long and 2 pairs of anterior setae 60 and 80 um long. Medial segment
with 2 pairs of setae 40 and 60 um long. Basal segment with 2 pairs of short setae each 24 um long and a
pair of longer setae each 68 um long. Anal ring slender, easily distorted, with a few minute pores and
6 setae each 88 um long. Circulus constricted, 130 um wide. Ostioles as in adult female, the anterior pair
often lying on ventral surface when mounted on slide but without setae. Anal lobes sclerotized on dorsal
and ventral surfaces, each lobe with a dorsal and ventral seta about 1-3 mm long, a ventral seta 0-8 mm
long and a dorsal 0-6 mm long. Ventral margins of third to eighth segments with sclerotized areas each
containing one long and one short seta, third and fourth segments with the longer seta 0-7 mm long,
44
Fig. 20 Malaicoccus moundi sp. n. Third instar female.
45
Fig. 21. Malaicoccus moundi sp. n. Second instar female.
46
Fig. 22. Malaicoccus moundi sp. n. Second instar male.
47
Fig. 23. Malaicoccus moundi sp. n. First instar.
48
fifth 0-6 mm, sixth 0-7 mm, seventh 0-6 mm and eighth 1:3 mm long. The short marginal seta on eighth
segment about 330 um long, the others on anterior segments shorter.
Body setae of different sizes. Dorsum with stout setae, most about 85 um long, in single transverse
rows but a median pair on last segment about 150 um long. A pair of sclerotized crescentic areas present
anterior to anal ring, each with a few setae of different sizes. Stout ventral setae also of different sizes,
a common length being about 100 um. Slender setae scattered on both surfaces, 8-28 um long. Other
minute setae numerous, each shorter than the diameter of its setal base. Trilocular pores not numerous.
Simple circular pores smaller than a trilocular pore, scattered.
Second Instar Female (Fig. 21)
This instar differs from previous instars in having shorter and fewer characters. Body about 1-25 mm long
and 0-85 mm wide. Head with 2 elongate sclerotized areas between antennae on dorsum. More extensive
sclerotization present around eyes and almost reaching anterior ostioles. Antennae 1-12 mm long; first
segment 130 um long, second 60 um, third 180 um, fourth 180 um, fifth 170 um and sixth 400 um. All
segments with numerous setae. Legs as in previous stage, hind trochanter+ femur 330 um long, hind
tibia + tarsus 350 um long. Claw 36 um long. Labium 128 um long and 78 um wide, with 14 pairs of setae.
Apical segment with 2 pairs of posterior setae, a pair of minute apical setae, 4 pairs of subapical setae
each 28 um long, a pair of lateral setae each 36 um long and 2 pairs of anterior setae 40 and 50 um
long. Medial segment with a single pair of setae 60 um long. Each lobe of basal segment with a pair of
short setae each 20 ym long and 1 long seta 72 um long. Circulus 80 um wide. Ostioles as in previous
stage, the anterior pair lying on ventral surface when mounted on slide, without setae. Anal ring with
1 or 2 minute pores and 6 setae each about 60 um long. Anal lobes each with dorsal and ventral elongate
areas of sclerotization, with 2 dorsal setae, one 1-0 mm long, the other 550 um long and 2 ventral setae,
one 1-0 mm long and the other 800 um long. Long marginal setae present on 6 preceding segments, each
setal base surrounded by sclerotized areas. Eighth segment with seta 1-0 mm long and a shorter about
180 um long, seventh with seta 300 um long, sixth seta 370 um long, fifth and fourth each 500 um and third
630 um long.
Dorsum with stout pointed setae in single transverse rows, usually 44-88 um long except in mid-region
of eighth segment where there is a pair 112 um long. Similar setae present in a sclerotized crescentic area
anterior-lateral to anal ring. Other dorsal setae more numerous and minute, each with seta much shorter
than diameter of base. Ventral surface with similar long setae to those on dorsum except on head margin
where there is a pair 100 um long and behind first ostioles where there is a pair each about 120 um long,
these on sclerotized bases. On abdomen there are shorter pointed setae each about 8 um long and minute
setae of two types. One of these is similar to the dorsal minute setae but the other type is longer 24 ym
but always shorter or same length as setal base. Trilocular pores on both surfaces, not numerous. Simple
circular pores present, sparse, smaller than trilocular pores.
Second Instar Male (Fig. 22)
Body elongate, head margin tending to be straight, 1:12 mm long and 0-65 mm wide. Antennae with
6 segments about 745 um long, the first segment 90 um long, second 45 um long, third 120 um, fourth and
fifth each 110 um long and sixth segment 270 um long. Fourth to sixth segments with slender setae
ranging in length from 36 to 52 um. Legs with hind trochanter + femur 280 um long, tibia + tarsus 300 ym
long. Claw 28 um long. Labium 120 ym long and 68 um wide, with 13 or 14 pairs of setae. Apical segment
with 2 pairs of posterior setae, a minute apical pair, 4 pairs of subapical setae each 20 um long, a lateral
pair each 36 um long, | or 2 pairs of anterior setae, the longer 64 um and the shorter when present 40 um.
Medial segment with 1 pair each 60 um long and basal segment with 2 pairs 18 um long and a pair 72 wm
long. Anal ring with a few minute pores and 6 setae. Anal lobes sclerotized on both surfaces, the dorsal
surface with 1 seta 0-9 mm long and another 500 um long, the ventral surface with one 0-9 mm long and
another 650 um long. Long marginal setae on 6 anterior segments with bases surrounded by sclerotized
areas, on eighth segment these setae 0-9 mm long, on seventh segment 220 um, on sixth 310 um, on fifth
400 um, on fourth 500 um and on third 550 um long.
Dorsal and ventral setae similar to those of second instar female but minute setae absent on venter
and sparse on dorsum. Trilocular pores few. Simple circular pores quite sparse, each smaller than a
trilocular pore. Multilocular disc pores each with 8 loculi, present on dorsum in small submedian groups
on fifth abdominal segment and anterior segments to prothorax, an occasional pore also present on
anterior abdominal segments and mid-thorax. Oral collar tubular ducts present, each slightly larger in
diameter than a trilocular pore and only slightly longer than wide with deep sclerotized internal collar.
On dorsum they tend to lie in single transverse rows but on head and thorax they become more scattered.
On venter they are present mainly on head and thorax and become sparse on abdomen.
49
First Instar (Fig. 23)
Body oval, 1-0 mm long and 0:65 mm wide, head sclerotized around eyes. Antennae 565-700 um long,
one of largest with first segment 80 um long, second 60 um, third 90 um, fourth 100 um, fifth 100 um and
sixth 270 um long. Each segment with a few long pointed setae at distal end but more numerous setae
present on last segment. Legs well developed with hind trochanter + femur 230-250 um long, hind tibia
+tarsus 250-270 um long. Claw 28 um long. Circulus apparently absent. Labium 92 ym long and 52 ym
wide, with 14 pairs of setae. Apical segment with 2 pairs of posterior setae, a minute pair of apical setae,
4 pairs of subapical setae each 20 um long, a lateral pair 38 um long and 2 pairs of anterior setae 20
and 56 um long. Medial segment with 1 pair of setae 52 um long and basal segment with 2 pairs 12 um
long and 1 pair 44 um long. Anal ring slender with a few minute pores and 6 setae each 52 um long.
Anal lobes sclerotized on both surfaces, each with 2 dorsal setae, one 1:0 mm long, the other 460 ym
long and 2 ventral setae, one 1-0 mm long and the other 700 um long. Long marginal setae present on
6 preceding segments, the bases surrounded by sclerotized areas. All of these setae stout, measuring on
eighth segment 1:0 mm long, seventh segment 160 um, sixth segment 220 um, fifth segment 280 ym,
fourth segment 280 um and on third segment 340 um.
Body setae 28-44 um long on dorsum but mid-region of eighth and last segment with pair of setae each
90 um long on sclerotized patches. Ventral setae usually longer, about 80 um long. Trilocular pores and
simple circular pores sparse.
Holotype 2, MALAYSIA: Malaya, Genting Highlands, 50 km E. of Kuala Lumpur, on stems of
Uncaria sp. [Naucleaceae], attended by the ant Hypoclinea sp., 28.ix.1973 (L. A. Mound) (BMNH,
London).
Not included in type-series. MALAYSIA: 2 third instar 9, 3 second instar 9, 3 second instar 3
and 9 first instars, same data as holotype (BMNH, London).
Discussion. The nearest species is M. khooi but it differs in the characters given in the discussion
of that species. It differs from M. riouwensis and M. takahashii in having 4 pairs of long marginal
setae on the abdomen instead of only 2 pairs. The short dorsal and ventral setae are similar to
those of M. riouwensis and M. takahashii in that the distance between the setal bases is greater
than the length of a seta but it differs from M. formicarii, which has the distance between the setal
bases shorter than the length of a seta.
Malaicoccus riouwensis Takahashi, 1950
Malaicoccus riouwensis Takahashi, 1950 : 66. Holotype, 2, INDONESIA: Riouw Is, Rempang, in association
with ants of genus Polyrhachis sp. (probably lost).
Adult Female (Fig. 24)
External appearance described originally as ‘Shining blackish brown, with no wax, in life, but dark
reddish brown and a little shining when dry.’
Body of adult female pear-shaped with a constriction opposite first coxae. Anal lobes moderately devel-
oped. Anterior end of body with extensive sclerotization, which on the dorsal surface, extends around the
antennal bases, then laterally and posteriorly around the first pair of ostioles. Ventrally the sclerotization
extends over anterior end of head to just above clypeus. Circulus folded in available specimen but described
by Takahashi as having deep constrictions at the sides and widest at anterior end. It is possible that
Takahashi studied specimens that were distorted and that in reality the circulus is widest at posterior end.
Labium 180 um long, with 21 pairs of setae. Apical segment with 2 pairs of posterior setae, a pair of
minute apical setae, 4 pairs of subapical setae each 28 um long, 1 pair of lateral setae each 60 um long and
2 pairs of anterior setae, the longest 80 um long. Basal segment with 2 pairs short setae and | pair longer
setae. Antennae 6-segmented, 2:33 mm long, thus longer than body; first segment 280 um long, the second
120 um, third 390 um, fourth 370 um, fifth 320 um and sixth 850 um and distinctly curved. Each segment
densely covered with short setae ranging from 24 um long on the proximal segments to about 40 um
towards terminal segment. Other slender setae 120 um long on second to sixth segments. Thick setae
present on all segments, these about the length of the shortest slender setae but much fewer. Legs well
developed, hind trochanter + femur 640 um long, hind tibia + tarsus 650 um long. Hind coxa with trans-
lucent pores on outer dorsal half. Short setae present on all leg segments, those on tibia numerous, about
28 um long, the distance between setal bases shorter than length of one seta. Claw 48 um long. Anal ring
slender with 6 setae about 56 um long and a few minute pores. Ostioles well developed, of the type com-
mon to the genus, the anterior pair with setae on the lower lips, the posterior pair without setae. Long
50
Le ent
i
a [ey ham
Lg,
et
ievu\a ©
he en
Fig. 24 Malaicoccus riouwensis Takahashi. Adult female. Specimen from Riouw Is, Rempang.
Antenna, hind leg and dorsal setae.
marginal setae on anal lobes and eighth segment only, the two preceding segments with much shorter
setae. Anal lobe setae comprising two long dorsal setae on an elongate area of sclerotization, the long
setae about 0-9 mm long and the short setae about 350 um long. The long ventral setae on eighth segment
0-8 mm long and a short seta 130 um long on an elongate-oval sclerotized area. Seventh segment with one
seta 220-250 um long and a shorter seta 72 um long on a round sclerotized area. Sixth segment with a
seta 350-400 um long and a short seta 72 um long also on a round sclerotized area.
Body setae of various lengths. On the dorsal surface of head some of the stout setae are 30-44 um long
with a few reaching 180 um. Others on posterior abdominal segments 44-100 um long, the longest on
mid-line. Stout setae on venter 40-68 um long on all segments especially in median areas. Some setae on
posterior abdominal segments 80 um long. Over the rest of the body there are abundant minute slender
setae 5-20 um long, the larger proportion being the smallest. The setal bases are spaced apart about 1-2
times the length of a single seta. Ventral surface with a distinct tuft of short setae opposite first spiracles,
the distance between these setal bases shorter than length of one seta. Trilocular pores present on both
surfaces, sparse except for a few between the first spiracles. Minute simple pores few, each with a diameter
smaller than a setal base. Tubular ducts represented by one or two on inner side of second and third coxae.
MATERIAL EXAMINED. INDONESIA: 1 9 (paratype), Riouw Is [Riau Is], Rempang, i.1946 (R. Taka-
hashi) (USNM, Washington).
Discussion. Mr Charles Chia-chu Tao has indicated that there is no type-material of this species
in the Taiwan Agricultural Research Institute, Taipeh. Dr Sadao Takagi of the Entomological
Institute, Hokkaido University, Sapporo, has informed the writer that there are no specimens in
Takahashi’s collection there and it must be assumed that the holotype is lost. Dr D. R. Miller
of the United States Department of Agriculture, Beltsville, has kindly made available a single
51
paratype from the collections of the United States National Museum. This specimen is not in
good condition and it has not been possible to illustrate it completely here.
The species is closely related to M. takahashii in having only 2 pairs of marginal setae nearly
1-0 mm long on abdomen on eighth segment and anal lobes. Anterior marginal setae to these
are only 200-400 um long. The most distinctive character separating this species from M.
takahashii is the arrangement of the setae on the hind tibiae. These are short, about 28 um
long, and the distance between the setal bases on the disc of tibia is shorter than the length of one
seta.
Malaicoccus takahashii sp. n.
Adult Female (Fig. 25)
An oval species, 1-7 mm long and 1-1 mm wide, with a deep constriction opposite first coxae. Anal lobes
moderately developed. Anterior end of body heavily sclerotized; on the dorsum this sclerotization en-
circles antennal base and curves laterally and posteriorly to completely surround anterior ostioles. The
sclerotization on ventral surface of head occupies the area immediately in front of clypeus to anterior
margin but curves forward laterally before reaching ostiole. Antennae with the first segment 250 um
long, second 150 um, third 360 um, fourth 360 um, fifth 350 um and sixth 720 um, the last segment curved.
With a total length of 2-19 mm the antennae are longer than body. All antennal segments densely covered
with short setae which range in length from 28 um on the first segment to 48 zm on the last segment.
Legs well developed, hind trochanter + femur 560 um long, hind tibia + tarsus 550 um long. All segments
with quite stout setae in moderate numbers, the smallest on tarsus about 44 um long with the space
between setal bases usually same or greater than length of an average seta. Hind coxa with translucent
pores on dorsal outer half. Claw 48 um long. Circulus notched at either side, 188 um wide. Labium
200 um long and 130 um wide with 20 pairs of setae, many of which are broken but their presence deter-
mined by setal bases. Apical segment with 2 pairs of posterior setae, a pair of minute apical setae, 4 pairs
of subapical setae, 1 pair of lateral setae and 2 pairs of anterior setae. Medial segment with 7 pairs of
setae and basal segment with 2 pairs of short setae and 1 pair of longer setae. Ostioles well developed, the
anterior pair each with sclerotized posterior lip containing a few setae. Anal ring rather thin and easily
distorted, with a few minute pores on periphery and 6 setae, each about 68 um long. Long marginal setae
each about 780 um long on anal lobes and eighth segment only. Dorsal surface of anal lobes with an oval
sclerotized area. Long marginal setae of eighth segment at apex of an elongate sclerotized area that also
contains a seta about 80 um long, each on a small sclerotized patch. Stout setae on dorsum confined to
head, prothorax, eighth and posterior segments. On the head they are present mainly within the sclero-
tized areas but on the anal lobe segment they are present in two sclerotized crescentic areas anterior-
lateral to anal ring. Although most of the stout setae are about 40 um long some on eighth segment may
reach 80 um and others on head attain a length of 100 um. Stout setae present on venter on all segments
but on abdomen they are present in median areas only. These setae are about 40 um long and on posterior
segments of abdomen they are up to 90 um long. Other body setae short, slender and numerous, evenly
distributed over dorsal and ventral surfaces but absent on dorsal parts of head and thorax. A marginal
tuft of short setae present opposite first spiracles, the distance between setal bases shorter than length of
one seta. All the short setae range in length from 5 to 24 um but the shorter setae are most numerous.
Trilocular pores few but there is a noticeable concentration between the two first spiracles. Tubular ducts
represented by one or two on inner side of second and third coxae. Simple circular pores minute, sparse.
Immature Instars
The three following instars are represented by poor distorted material and it has not been
possible to illustrate them. Nevertheless, the long marginal setae are present only on the anal
lobes as in the adult.
Third Instar Female
Body oval 1:25 mm long and 0-8 mm wide, constricted opposite first coxae. Head sclerotized around
antennal bases, between clypeus and anterior head margin and around eyes on venter. Antennae 1-55 mm
long, first segment 180 um long, second segment 100 um, third 250 um, fourth 250 um, fifth 200 um and
sixth 570 um long; all segments with numerous slender setae. Legs with hind trochanter + femur 400 um
long, hind tibia+tarsus 410 um long. Claw 40 um long. Circulus 128 um wide. Anal ring with a few
52
SN
Fig. 25. Malaicoccus takahashii sp. n. Adult female. Specimen from Malaya, Kuala Lumpur.
3
minute pores and 6 setae each 60 um long. Labium 160 um long and 90 um wide, with 16 pairs of setae.
Apical segment with 2 pairs of posterior setae, a pair of minute apical setae, 4 pairs of subapical setae
each 20 um long, | pair of lateral setae each 56 um long and 2 pairs of anterior setae, the longest 64 um
long. Medial segment with 3 pairs of setae, the longest 56 um long. Each lobe of basal segment with a
pair of short setae 20 um long and one seta 44 um long. Anal lobes sclerotized, each with one dorsal
seta 0-8 mm long and a shorter seta 120 um long, the ventral surface with one seta 0-8 mm long and another
280 um long. Eighth segment with marginal setae 0-7 mm long and anterior marginal setae much shorter,
each about 100 um long, all on sclerotized areas. Dorsal and ventral setae numerous, of two types, a
slender pointed type and a minute type with the seta much shorter than diameter of setal base.
Second Instar Female
Body 1-0 mm long and 0-55 mm wide, sclerotized on head between antennae on dorsum and around eyes
on venter. Antennae 1-04 mm long, first segment 120 um long, second 60 um, third 170 um, fourth and
fifth each 160 um and sixth 370 um long. All segments with numerous slender setae. Hind trochanter +
femur 290 um long, hind tibia+tarsus 320 um long. Claw 34 um long. Circulus 80 um wide. Anal ring
slender with a few minute pores and each of the 6 setae 48 um long. Labium 130 um long and 70 um
wide, with 14 pairs of setae. Apical segment with 2 pairs of posterior setae, a pair of minute apical setae,
4 pairs of subapical setae each 20 um long, | pair of lateral setae 40 um long and 2 pairs of anterior setae
the longest 56 um. Medial segment with a single pair of setae 44 um long. Each lobe of basal segment
with a pair of short setae 16 um long and one seta 36 um long. Anal lobes each with 4 setae, a dorsal and
ventral 0-8 mm long, one dorsal 150 um long and one ventral seta 270-320 um long. Eighth segment with
marginal setae 0-8 mm long. Anterior marginal setae 100 um long on sclerotized areas. Trilocular pores,
simple circular pores and minute setae very few.
First Instar
Body oval, 0-85 mm long and 0:45 mm wide. Antennae 680 um long, the first segment 70 um long, second
40 um, third 100 um, fourth 110 um, fifth 100 um and sixth 260 um long. Hind trochanter + femur 220 um
long, hind tibia+tarsus 240 um long. Claw 28 um long. Anal ring setae 40 um long. Anal lobes each
with a dorsal and apical seta 0-85 mm long and a short dorsal and ventral seta 220 um long. Eighth seg-
ment with marginal setae 0-85 mm long and anterior marginal setae 100-120 um long, all on sclerotized
areas. Labium 110 um long and 60 um wide with 14 pairs of setae. Apical segment with a pair of minute
apical setae, 4 pairs of subapical setae each 20 um long, a pair of lateral setae 32 um long and 2 pairs of
anterior setae, the longest 60 um long. Medial segment with 1 pair of setae 52 um long and each lobe of
basal segment with a pair of short setae 16 um long and 1| seta 40 um long.
Holotype 2, MALaysIA: Malaya, Selangor, Kuala Lumpur, 11.viii.1943 (R. Takahashi) (BMNH,
London).
Not included in type-series. MALAYSIA: 1 third instar 9, 9 second instar 9 and 2 first instars,
same data as holotype (UM, Kuala Lumpur).
Discussion. This is probably the undescribed species from Kuala Lumpur, on Palaquium obova-
tum (Sapotaceae), mentioned by Takahashi (1951) when discussing M. formicarii. Specimens
were kindly made available by Dr Khoo Soo Ghee.
The most distinctive features of the adult female are the long pairs of setae present only on the
anal lobes and eighth segment. These are about 780 um long. The only other marginal setae, in
addition to the normal body setae, are present on the seventh segment but these are only 80 um
long. A major character separating this species from M. riouwensis is the arrangement of the tibial
setae, which although longer than those in M. riouwensis, are sparser, and on the tibial disc the
distances between the setal bases are the same as or longer than the length of one seta.
PARAMYRMOCOCCUS Takahashi, 1941
Paramyrmococcus Takahashi, 1941 : 204. Type-species: Paramyrmococcus chiengraiensis Takahashi, by
original designation and monotypy.
DEscrIPTION. Adult female with turbinate to oval body, tapering to a rounded posterior end, anal lobes
obsolete. Antennae 6-segmented, about as long as body, first and second segments each with 1 or 2 stout
setae, third to sixth segments densely covered with slender setae; second segment quite small, either
shorter or just slightly longer than wide. Legs well developed, setae varying in size and number, claw
54
Nth
it th li
tlt! 4 iiss
ve
(1) Wy
Hy! A i!
4
nn ni NN S B
ae Lhe
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\S
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UGA BE ifn
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by WO A TaN f
Firing” FPN Mig ihe, Vi ih Vy; NEATH \ AW
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ij Ni
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aM ili ii Lin Lanne Nay
RAL)
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ba CN ey
bilyrag 7 \ wi, iH VEZ BGS Ww \\ i
Nt ee 16 ep SEs ZAR xe Ae Sy my
NU ne ag Zhi VUES SS a Br SRA ‘i
‘
, MESSRS ‘i epi Vt
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iA iy ay Hy Wyn te a Nii, iy Te
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VY
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wiht LA aA se iN Ais s
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i},
ut MN, DJ.w
qi aly ae <
ies
Fig. 26 Paramyrmococcus chiengraiensis Takahashi. Adult female. Specimen from Thailand,
Chiengrai.
a3
stout with a pair of wide flat digitules. Anal ring sclerotized with 6 setae but without pores. Ostioles
situated on margins, of unusual development with large prominent lips that are heavily sclerotized but
are devoid of setae. Circulus normal, usually oval. Body setae minute and slender, of unusual abundance,
covering almost entire surface except for intersegmental areas, lying in definite directions giving the
surface a wavy appearance. Other setae stout, situated mainly around margins; anal lobe setae not much
more than twice diameter of anal ring in length. Trilocular pores few.
DIscussION. This genus is easily separated from the others in the Allomyrmococcini in being
without recognizable anal lobes and in having short setae, only about twice diameter of anal ring
in length, in normal positions of lobes. The immature instars are about the same shape as the
adult but they tend to be more slender. Distinctive characters of the immature instars, based on
P. vietnamensis, are short cylindrical and blunt setae on dorsum at least, which are replaced
completely in the adult by short pointed setae.
DISTRIBUTION. The genus is known so far from Thailand and Vietnam.
Key to species of Paramyrmococcus (adult females)
Setae on hind tibial disc sparse, the distance between setal bases longer than length of a single seta.
Marginal stout setae on head and thorax attaining a length of 40 um_ chiengraiensis Takahashi (p. 56)
Setae on hind tibial disc numerous, the distance between setal bases shorter than length of one seta.
Marginal stout setae on head and thorax attaining a length of 80 um vietnamensis sp. n. (p. 61)
Paramyrmococcus chiengraiensis Takahashi, 1941
(Fig. 26)
Paramyrmococcus chiengraiensis Takahashi, 1941 : 204. Holotype 2, THAILAND: Chiengrai, Lampang,
on undetermined vine, attended by ant Dolichoderus sp. [= Hypoclinea sp.] (probably lost).
Paramyrmococcus chiengraiensis Takahashi; Takahashi, 1942: 16.
Adult Female (Fig. 26)
External appearance in life not noted. On the slide, adult female turbinate or broadly oval, widest at
mesothorax, posterior end of body tapering, rounded, about 2-0 mm long and 1-25 mm wide. Antennae
about 1-78 mm long, first segment 170 um, second 90 um, third 430 um, fourth 320 um, fifth 270 um and
sixth 500 um long. Second segment about 80 um wide and thus slightly longer than wide. Sixth segment
widest at proximal half, which then narrows before becoming slightly swollen. The first two segments
with 1 or 2 stout setae only, the remaining segments densely covered with slender setae about 32 um long
on third segment and as long as 44 um on sixth segment. Legs well developed, hind trochanter + femur
520 um long, femur at its widest about 150 um; hind tibia+ tarsus 450-460 um long, the tarsus 80 ym
at its widest and distinctly shorter than trochanter + femur. Slender setae on dorsal and ventral surfaces
of tibia quite sparse, about 40 um long, the spaces between setal bases much greater than length of one
seta. Claw 40 um long. Labium 160 um long and 140 um wide, with 14 pairs of setae. Apical segment
with two pairs of posterior setae, a pair of minute apical setae and four pairs of subapical setae each
28 um long except the second pair, which are more slender and shorter, 20 um long. Lateral setae and 2
pairs of anterior setae present, each 40 um long. Medial segment with 1 pair of setae each 40 um long
and each lobe of basal segment with 2 pairs of setae 20 um long and 1 pair 40 um long. Circulus oval to
subrectangular, 60-104 um wide. Ostioles well developed, on the margins, with protruding sclerotized
lips, without setae. Anal ring 72-80 um wide with 6 setae 70-80 um long.
Body setae mainly slender, about 16 um long but with a smaller proportion of longer setae about
20 um long evenly distributed among the smaller, covering almost entire surface except for intersegmental
areas; the distance between the setal bases much shorter than the length of one seta. On the dorsum the
setae are even closer together than on venter. Other setae present are stout, those on margins of head and
thorax on both surfaces 40-50 um long, on margins of abdomen 60-80 um long except on eighth segment
where they are 112 um long and the two ventral anal lobe setae about 120 um long. Mid-ventral setae
about 40 um long. Trilocular pores sparse, apparently on venter only. Round pores on dorsal margins
few, each slightly smaller than a trilocular pore and with obscure structure.
MATERIAL EXAMINED. THAILAND: 2 9, ‘Siam’, Chiengrai, 16.iv.1940 (R. Takahashi) (TARI,
Taipeh).
56
Ne
ANS Zo)
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rine
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NH CE
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HENNY ANNE Ps WWF
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an
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RZ its HN Ne
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NS
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es ir
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= \
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ifs
ag
NN
Fig. 27 Paramyrmococcus vietnamensis sp. n. Adult female. Specimen from Vietnam, Da Lat.
a7
TPN
i Pal Avy
oe ARNO
VN YT PULLIN AS
tie 1 PN
= ‘ )
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re |
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; Aad] J ie
/ tee
Fig. 28 Paramyrmococcus vietnamensis sp. n. Third instar female.
58
Fig. 29. Paramyrmococcus vietnamensis sp. n. Second instar female.
Ros
ey \ .
ee
Fig. 30 Paramyrmococcus vietnamensis sp. n. First instar.
| i D.J.W.
Discussion. The two specimens examined seem to belong to Takahashi’s original type-series.
They were kindly made available by Mr Charles Chia-chu Tao of the Taiwan Agricultural
Research Institute, Taipeh.
Paramyrmococcus vietnamensis sp. n.
Adult Female (Fig. 27)
External appearance not known. When mounted on the slide, body turbinate, widest at mesothorax,
posterior end narrow and rounded, length 2:1 mm, width 1-3 mm. Antennae 1-9-2-0 mm long and thus
about as long as body. On an antenna measuring 1-9 mm long the segment lengths are: first 200 um,
second 60 um, third 470 um, fourth 290 um, fifth 290 um and sixth 590 um. The second segment 80 um
wide and, therefore, wider than long; sixth segment with a constriction at about middle but basal part
wider than terminal part. First and second segments each with one or two stout setae only, remaining
segments densely covered with slender setae ranging from about 40 um long on third segment to 60 um
long on sixth segment. Legs well developed, hind trochanter + femur 560-570 um long, the widest part of
femur about 150 um at distal end; hind tibia + tarsus 560-590 um long, thus same length as trochanter +
femur or longer, widest part of tarsus 80 um. Legs with 1 or 2 stout blunt setae and covered with slender
setae about 40 um long, which on the disc of hind tibia are set close together on dorsal and ventral surfaces
so that spaces between are shorter than length of one seta. Claw 40 um long. Ostioles well developed,
situated on margins, each with prominent heavily sclerotized lips without setae. Circulus oval, 56-60 um
wide but sometimes with slight constrictions on either side, situated between fourth and fifth segments.
Labium 172 um long and 144 um wide with 14 pairs of setae. Apical segment with 2 pairs of posterior
setae, a pair of minute apical setae, 4 pairs of subapical setae each 28 um long, | pair of lateral setae 40 um
long and 2 pairs of anterior setae, the longest 60 um. Medial segment with 1 pair of setae each 44 um
long and basal segment with 2 pairs of setae 24 um long and 1 pair 40 um long. Anal ring 68-72 um wide,
with 6 setae about 80 um long. Body setae abundant and covering almost entire surface except for inter-
segmental spaces. For the most part the setae are slender, about 16 um long on dorsum and 16-20 ym
long on venter, the spaces between the setal bases much shorter than length of one seta but even so these
spaces are shorter on dorsum than on venter. Some of the slender setae are a little longer and are easily
discernible by the larger setal bases. The directions of the slender setae form definite patterns giving the
surface a wavy appearance. Stout setae, easily recognized by the large setal bases, present in small numbers
around margins of both surfaces and across ventral abdominal segments, about 25-40 um long but on
the normal positions of the lobes there are usually 2 dorsal and 2 ventral setae about 120 um long and on
margins of eighth segment there are 1 or 2 setae about 100 um long. Trilocular pores apparently confined
to venter where they are sparse. Dorsal marginal and submarginal areas with a few round pores slightly
smaller in diameter than a trilocular pore but with an obscure structure.
Third Instar Female (Fig. 28)
Body pyriform, 1-7 mm long and 0-7 mm wide. Antennae 1:57 mm long, the sixth segment with a con-
striction in the middle. First and second segments with a few stout setae only, third to sixth segments
with numerous slender pointed setae but not so numerous as in adult. Labium 160 um long and 120 ym
wide with same number of setae as in adult. Apical segment with 4 pairs of subapical setae each 24 ym
long, 1 pair of lateral setae each 32 um long and 2 pairs of anterior setae 40 um long. Medial setae 40 um
long and basal setae 20 zm and 36 um long. Legs with hind trochanter + femur 450 um long, hind tibia +
tarsus 460 um long. Apart from stout setae 40-55 um long there are also numerous slender pointed setae
28-32 um long. Claw 36 um long. Ostioles prominent with wide sclerotized lips. Circulus about 60 um
wide with a distinct notch at either side. Anal ring 64 um wide with the 6 setae about 72 um long.
Dorsal surface with numerous setae of different sizes. Long stout marginal setae present about 60 um
long, each usually with a blunt or irregular apex and often showing a discharge. Shorter pointed setae
on margins of head and thorax and on last segment. Evenly distributed over dorsum are two types of
minute setae, blunt with parallel sides. One type is extremely thin and about 6 um long, the other type
about twice as thick, about 5 um long and interspersed with the thin type, both types being present in
about equal numbers. Ventral setae only occasionally represented by the minute blunt type, replaced by
numerous pointed setae 16-20 um long, these also accompanied with long thick blunt setae. Trilocular
pores few, on ventral surface only but they are more numerous in median areas of thorax. Simple circular
pores, sparse, around dorsal margins.
61
Second Instar Female (Fig. 29)
Similar to third instar but with the following major differences. Body 1-4 mm long and 0:67-0:77 mm wide.
Antennae 1:05-1:10 mm long, with long stout setae on first two segments and much more slender setae
on third to sixth segments, these quite abundant and ranging in length from 35 to 50 um. Legs well
developed with hind trochanter + femur 320-350 um long, tibia + tarsus 320-350 um long. Claw 30-36 um
long. Labium 128 um long and 88 um wide, with setae similar to those of third stage but longest basal
setae only 32 um long. Circulus with a small notch on either side, about 36 um wide. Anal ring 52 um
wide, the anal ring setae about 60 um long. Body setae less numerous than in previous stage but with the
same two types of minute setae on dorsum. The same two types are present also on venter but they are
interspersed with long blunt setae and normal pointed setae. Trilocular pores sparse, confined to venter.
Simple circular pores present in small numbers around dorsal margins and on prothorax at a point
opposite clypeus.
First Instar (Fig. 30)
Body elongate-pyriform, about 1-2 mm long and 0:57 mm wide at thorax, posterior end of body rounded,
anal lobes barely perceptible. Antennae 720 um long, the first four segments each with a few stout setae
about 50 um long, the tips often damaged and each seta having a fleshy appearance. The two terminal
segments with much more numerous slender pointed setae, these also 50 um long; sixth segment with a
slight constriction near middle. Legs well developed, hind trochanter + femur 250 um long, hind tibia +
tarsus 260 um long. Claw 30 um long. Circulus situated between fourth and fifth segments, small and
oval, about 28 um wide. Anal ring 40 um wide, without pores but with 6 setae each about 45 um long.
Labium 108 um long and 80 um wide, differing from previous stage in that some setae are shorter, the
anterior setae 36 um long, medial setae 36 um long and basal segment with 2 pairs of setae each 12 um
long and 1 pair 32 um long. Ostioles on margins with heavily sclerotized lips, without setae.
Body setae of various sizes. Anal lobe setae comprising 3 dorsal and 2 ventral, each about 110 um
long, the bases of the ventral setae at the end of an elongate bar-like area of sclerotization. Margins of
dorsum with thick setae usually about 40 um long, each of which is irregularly blunt at apex and showing
what appears to be a discharge. Dorsum with thin type of minute setae only, about 6 um long. Anal lobe
segment without these setae but with pointed setae about 35 um long. Ventral surface with long setae
around margins, some of which attain a length of 50 um. Other long setae present in median areas of
segments, these interspersed with minute setae similar to those on dorsum accompanied with short
pointed setae 12-24 um long. Trilocular pores quite sparse, there being a few around the ventral margins
and on thorax.
Holotype 9, VIETNAM: South, Da Lat (37km SE.), Chute de Gauhgah, 25.iv.1960 (R. E.
Leech).
Paratypes and many immatures, VIETNAM: South, Da Lat, same data as holotype (16 9, BMNH,
London; 2 9, USNM, Washington; 2 2, TARI, Taipeh; 2 9, IEAUN, Portici; 2 9, RNH, Leiden).
DIsCussION. This species is close to P. chiengraiensis but differs mainly in the characters given
already in the key to species.
RHIZOECINAE
The genera Eumyrmococcus and Xenococcus are aberrant and are assigned to this subfamily
mainly on the bases of the width—length ratio of the labium and the long slender claws with short
setose digitules, characters which have been discussed in the section on classification.
Both genera and the South American genus Chavesia differ from others in the Rhizoecinae in
possessing minute setae densely covering the dorsum. Chavesia and Eumyrmococcus have the
cephalothorax strongly dilated and all three genera have the abdomen tapering or abruptly
narrowed. The three genera may be separated by the following key.
Key to adult females of the anomalous ant-attended genera of Rhizoecinae
1 Antennae 2-segmented, not more than 180 um long. Circulus absent EUMYRMOCOCCUS ao =
— Antennae 4- or 5-segmented, at least 280 um long. One or two circuli present
2 Antennae well developed, as long as body. Anal ring terminal. Two circuli present
XENOCOCCUS (p. 63)
— Antennae shorter than width of body. Anal ring situated some distance from apex of abdomen,
near base of well-developed anal lobes. One circulus present . ‘ : ; CHAVESIA
62
EUMYRMOCOCCUS Silvestri, 1926
Eumyrmococcus Silvestri, 1926: 271. Type-species: Eumyrmococcus smithii Silvestri, by original desig-
nation and monotypy.
Eumyrmococcus Silvestri; Williams, 1969 : 336.
Eumyrmococcus Silvestri; Williams, 1970: 138.
The description of this genus is virtually the same as the type-species given already by Williams
(1970). The genus was placed in the tribe Rhizoecini by Williams (1969) and this assignment has
been confirmed here by a further study of the labium which has a width-length ratio of about 1 : 2.
Eumyrmococcus smithii Silvestri, 1926
(Fig. 31)
Eumyrmococcus smithii Silvestri, 1926 : 273. LECTOTYPE 92, Cuina: Macao [on roots of plants, attended
by ant Acropyga (Rhizomyrma) sauteri Forel] (IEAUN, Portici), here designated [examined].
Eumyrmococcus smithii Silvestri; Silvestri, 1927 : 254.
Eumyrmococcus smithii Silvestri; Takahashi, 1934: 2.
Eumyrmococcus smithii Silvestri; Williams, 1970: 138.
Williams (1970) has described and illustrated this species recently and the illustration from the
Bulletin of Entomological Research is reproduced with permission but with slight modifications.
The labium was too distorted in the material studied at the time but further specimens of the type-
series have become available through the kindness of Dr A. Tranfaglia and it is now possible to
modify the description of the labium. There is no specimen with a perfect labium and the follow-
ing description of this structure remains tentative.
Length 156-160 um and 80 um wide, apical segment with 2 pairs of posterior setae, 1 pair of minute
apical setae, 4 pairs of subapical setae. One pair of lateral setae and 2 pairs of anterior setae. The sub-
apical setae become longer and stouter towards the base. Medial segment with 1 pair of medial setae and
basal segment with 2 pairs of short setae and 1 longer pair. Almost all the setae are curled in the specimens
examined and the lengths are difficult to determine.
MATERIAL EXAMINED. Eumyrmococcus smithii Silvestri, lectotype 9, CHINA: Macao (IEAUN,
Portici).
CHINA: 4 9, same data as lectotype (IEAUN, Portici) (1 9, BMNH, London); | 2, Shanghai
(IEAUN, Portici); 4 9, Taipo Market, 26.xii.1924 (F. Silvestri) (2 2, BMNH, London; 2 9,
IEAUN, Portici) (mounted from Silvestri’s alcohol material). TAIWAN: 4 92, Taichu, on sugar-
cane, 24.11.1933 (M. Yangihara) (TARI, Taipeh).
Discussion. The width-length ratio of 1:2 of the labium confirms the assignment of the genus
to the Rhizoecinae agreeing also to the definition of the subfamily by Koteja (1974a).
XENOCOCCUS Silvestri, 1924
Xenococcus Silvestri, 1924 : 312. Type-species: Xenococcus annandalei Silvestri, by original designation
and monotypy.
Xenococcus Silvestri; Silvestri, 1926 : 275.
Xenococcus Silvestri; Silvestri, 1927 : 253.
DESCRIPTION. Mealybugs with broadly-oval body, abdomen tapering to narrow sclerotized terminal
segment. Apex with dorsal anal ring projecting beyond ventral lobes. Anal ring with 8 setae, the anterior
2 pairs slender and the posterior pair thick and long, the posteriormost often on ventral surface. Anal
lobes with numerous long stout setae. Antennae 4-segmented, nearly as long as body, the first segment
long and wide, articulating with much narrower second segment containing small teeth at proximal
end fitting into grooves at distal end of first segment. All segments with long pointed setae. Legs long
and well developed with long slender claw, at base of which are 2 short setose digitules. Labium long and
slender with a width-length ratio in excess of 1 : 2. Subapical setae reaching more than half distance to
base of apical segment. Two circuli present in adult, circular and slightly conical. Body setae densely
covering dorsum. Ventral setae longer and not so numerous. Pores and ducts absent.
Immature instars similar in shape to adult but with fewer setae and with only a single circulus.
63
Z
7 nay
pnp PPP OG I,
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Gs -
hy? LG, a AG 75}
oc;
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/
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2
oe
é
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ARAL AS As! EE WA
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iWalii
V\-149
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IN
D.J.W.
Fig. 31. Eumyrmococcus smithii Silvestri. Adult female. Specimen from Japan.
64
Bi
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hi Y wi ah vin ya a ANNs x
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.
Fig. 32 Xenococcus annandalei Silvestri. Adult female. Specimen from India, Barkuda I.
65
DISCUSSION. The most distinctive character of this genus is the peculiar pair of antennae which
are unlike any known in the Pseudococcidae. An anal ring projecting beyond ventral anal lobes,
the long stout anal lobe setae and the dense covering of dorsal pointed setae, are characters shared
with some of the Allomyrmococcini but the long slender claw with short setose digitules, the
round sclerotized circuli and the 4-segmented antennae are characters which seem to belong to
the Rhizoecini of the subfamily Rhizoecinae.
De Lotto (1977) has recently described Xenococcus scorpioides, an interesting species from South
Africa, from the nests of Acropyga sp. and the first anomalous species from the African
continent. In possessing short 4-segmented antennae, shorter legs with thick claws, it does not
seem to be congeneric with X. annandalei but probably lies somewhere between Chavesia and
Xenococcus.
DISTRIBUTION. At present the genus is known from India, Malaya, Vietnam and Hong Kong.
Xenococcus annandalei Silvestri, 1924
Xenococcus annandalei Silvestri, 1924: 312. LECTOTYPE 9, INp1A: Barkuda I. [Chilka Lake, Madras
District (now Orissa Province), on roots of Ficus obtusa, with ants Acropyga acutiventris Roger]
(IEAUN, Portici), here designated [examined].
Xenococcus annandalei Silvestri; Silvestri, 1926 : 275.
Xenococcus annandalei Silvestri; Silvestri, 1927 : 253.
Adult Female (Fig. 32)
Adult female in alcohol ovoid, abdomen tapering, posterior end curved slightly upwards, antennae
strongly geniculate, lying in a posterior direction almost parallel to body margins.
On the slide, when flattened, broadly oval, 1-6-1-7 mm long and 1-1—-1-4 mm wide, abdomen tapering
to a narrow sclerotized last segment almost as long as broad with the anal ring at apex of dorsum pro-
jecting beyond a pair of ventral lobes that curve inwards almost to mid-line. Antennae with 4 segments,
of a distinctive shape and structure, 1:35-1:54 mm long, attached to dorsal surface of head. One of longest
antennae with first segment 380 um long and 230 um at widest. Remainder of antenna much narrower,
the second segment 440 um long, the third 200 um long, and last segment tapering, 520 um long. All
segments with numerous long slender setae of different sizes, the longest on third and fourth segments.
Length of longest seta, on third segment 500 um. Joint between first and second segments distinctive,
there being small tooth-like projections on proximal lower corner of second segment which fit into small
grooves on posterior distal edge of first segment. Legs well developed, hind trochanter+femur 410-
470 um long, hind tibia + tarsus 470-550 um long. Tarsus tapering to a distance of about half its length
which then becomes narrow and subparallel towards distal end, a small notch also present on inner
margin at point between tapering part and narrow extremity. Each leg segment with setae of various
lengths but mostly long and slender, the longest on femur about 180 um long. Claw unusually long and
narrow with a length of 40-44 um and with a pair of setose digitules, each 16 um long. Two circuli present
situated in middle of third and fourth segments. Each circulus round, 40 um in diameter, with a sclerotized
outer rim and with the inner surface forming a cup. Labium unusually elongate; in alcohol specimens
250 um long and 110 um wide but, when flattened on slide, about 128 um wide with 12 or 13 pairs of setae.
Apical segment tending to be lanceolate instead of the usual conical shape, with 2 pairs of posterior setae,
a pair of apical setae 20 um long, 4 pairs of subapical setae extending towards base of segment, the distal
setae about 20 um long and the others becoming longer towards the base. Anterior setae represented by a
single pair each 100 um long but a single lateral pair each 80 um long may be present or absent. Medial
segment with a single pair of medial setae each about 75 um long. Basal segment with a pair of setae each
10 um long and a single setae 64 um long on each lobe. Anal ring situated mainly on dorsal surface but
often curving to ventral surface, sclerotized, about 120 um wide at widest point, without pores but
apparently with 4 pairs of setae around perimeter, there being 2 pairs of slender anterior setae each about
110 um long and an outer pair of stouter setae each 400 um long. A pair of stout setae about 0-8 mm long
usually on ventral surface seem to belong to ring. Vulva occupying almost total width of its segment.
Posterior setae stout, each about 1-0 mm long, there being 4 such pairs, 1 pair on lateral edges of dorsum
anterior to anal ring and 3 pairs on posterior edge of ventral lobes. An anterior pair of setae on inner
edges of these lobes much shorter and more slender.
Dorsal setae abundant, covering almost entire surface except for head and intersegmental areas, the
spaces between setal bases much shorter than length of one seta. The most abundant are slender, 20-28 um
long but there are some, interspersed with these, about 40 um long and which may be detected by the
66
iis
MAYS \
Waly
ii
it
Li
1A
i “4 APA!
ARAN
ee alg
Minh Ay titi
itl y Wilis
LR lk
ny ety Hi tthays ats
ii
TN " 4 TU ev
SAR HU amr AILS eC We et a i
Hit WTEC eg eth itd li iil
» (Gy etek} WAN as 1 =
\, th Mente A, wth Ay atta
4 AARY thd %
CREA Se A SP E
\\
\ ANS Vtg Why Wye 4 NG :
Vat WH a a A
Naess Wari Sas yh ne
AR Pain i, Neh ke =
. \
eh
vil =e
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reenz Pan) is
Nias! ji \
Wy CLLEL L
Tish
ay TAINAN
\\
j Fe (he
if \ i
Fig. 33 Xenococcus annandalei Silvestri. Third instar female.
67
~“. ¥
KAS |
fe alps
i}
Wh oss
|
\ f 2
A ree =
\ \\ NAN
Poo NC \
fr | 4
VES ; ~f)
i] Hats \ 11 \
Ls ut \\ ]
Wm \
NS \ AA Vow a
\ ey x \
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>] / | |
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Fig. 34 Xenococcus annandalei Silvestri. First instar. Specimen from India, Mysore, on
coconut roots.
68
slightly larger setal bases. On prothorax and mesothorax there are short flat setae in varying numbers
but usually few. Each is variously shaped and often peculiarly angled. Setae on last segment longer than
usual body setae and just anterior to anal ring a fringe of long setae present comprising 2-3 median pairs
each about 480 um long and a lateral pair usually stouter about 560 um long. Other long setae, some
attaining a length of 240 um, present on head and around margins of abdomen. Ventral setae numerous,
of different sizes, but many are long, the longest about 200 um. Minute setae present from about fifth
segment as far forward as prothorax in a zone extending to spiracles. Body pores and tubular ducts
absent.
Third Instar Female (Fig. 33)
Body ovoid, 1:15 mm long, tapering to last segment. Posterior end of body rounded, last segment sclero-
tized at least on dorsum, the ventral surface curved inwards to form a pair of lobes. Antennae with 4 seg-
ments, 1:10-1:23 mm long; on longest antenna, first segment 350 um long, second 360 um, third 180 umand
fourth 340 um, narrower than other segments. Joint between first and second segments similar to that in
adult female and with minute teeth on outer posterior angle of second segment fitting into small grooves
on outer distal angle of first segment. Each segment with long slender setae of different lengths. Legs
similar in shape to those in adult female but shorter, hind trochanter+ femur 280-310 um long, hind
tibia+ tarsus 350-400 um long. Labium as in adult but 205 um long and 88 um wide, with 13 pairs of
setae. Apical segment with 2 pairs of posterior setae, a pair of minute apical setae 12 um long, 4 pairs of
subapical setae set wide apart so that proximal pair at mid-point of segment, the setae ranging in length
from 12 um at apex to 68 um on mid-segment. Also present 1 pair of lateral setae and 1 pair of anterior
setae each 68 um long. Medial segment with 1 pair of setae 60 um long. Basal segment with each lobe
containing 2 short setae each 8 um long and | seta 36 um long. One circulus present, circular, about
32 um in diameter, situated in middle of fourth segment. Anal ring at apex of dorsal surface about 56 um
wide, with narrow sclerotized rim and surrounded by 4 pairs of setae. The anterior median pair are slender
and only slightly longer than diameter of ring, outer 2 pairs stouter, the third pair about 210 um long and
the fourth pair 440 um long. Anal lobes each with one dorsal and two ventral stout setae about 1-0 mm
long, there being also a slender seta about 150 um long on inner margin of lobe.
Dorsal setae abundant anterior to fourth abdominal segment, these mainly slender 12-16 um long,
the distance between setal bases usually shorter than length of one seta or about same length. Minute
flat setae with varying distal shape, scattered among slender setae on prothorax and mesothorax. Dorsal
setae on fourth and posterior segments fewer and of different sizes, the longest about 180 um. Ventral
setae mainly long and slender on abdomen, the longest about 160 um but from first segment to prothorax
small slender setae, similar to those on anterior dorsum extend inwards from margin to spiracles. Pores
and tubular ducts absent.
First Instar (Fig. 34)
Body oval, 0-7-0-9 mm long, membranous except for anal lobe segment which on dorsum is sclerotized
on margin extending to a band anterior to anal ring. Anal lobes present ventrally, these also sclerotized.
Antennae with 4 segments similar in shape to those of previous stages, 0:85-0:90 mm long, the longest
antenna with first segment 240 um long, the second 320 um, third 110 um and fourth 230 um long. Each
segment with slender setae varying in length, the longest about 300 um. Joint between first and second
segments similar to those in other stages. Legs as in previous stages except that tarsus longer than tibia
whereas in other stages the tibia is longer than tarsus. Hind trochanter + femur about 220 um long, hind
tibia+ tarsus 240-260 um long. Claw slender, 28 um long. Labium 180 um long and 80 um wide with
similar setal distribution to that in previous stages but setae longer than in third stage. Apical segment
with a pair of apical setae each 8 um long, 4 pairs of subapical setae extending to a point more than half
length of segment and ranging ‘a length from 12 um distally to an inner pair 76 um long. Also present are
a lateral pair and an anterior pair each 76 um long. Medial segment with either | or 2 pairs of setae, the
longest 72 um. Basal segment with a pair of setae each 8 um long and | seta 72 um long on each lobe.
Circulus round, about 28 um in diameter, situated within borders of fourth segment. Anal ring entirely on
dorsum, with narrow rim and 4 pairs of setae around outer margin, there being 2 pairs of inner slender
setae each about 60 um long, a third stouter pair about 360 um long and a stout posterior pair about
750 um long. The last setae resemble anal lobe setae which are of similar shape and length, there being
on each lobe, a pair on dorsal margins and a single seta at apex of lobe on venter.
Dorsal setae mainly short and slender on anterior abdominal segments, thorax and head, each about
12 um long. A few flat and irregularly shaped setae present on prothorax each about 8 um long. Ab-
dominal setae all slender, of different lengths, the longest always in transverse rows, about 120 um long.
69
Ventral setae on abdomen similar to those on dorsum of abdomen but fewer. Setae sparse on thorax
and head. Pores and tubular ducts absent.
MATERIAL EXAMINED. Xenococcus annandalei Silvestri, lectotype 9, INDIA: Barkuda I. (IEAUN,
Portici).
INDIA: 7 9, same data as lectotype; 2 9, 1.x.1922 (IEAUN, Portici), 1 9 and 1 third stage 9
(BMNH, London), | 9 and | third stage 9(USNM, Washington); 8 9, on Ficus religiosa, 28.vi.1922
(N. Annandale) (7 9, on slides and in alcohol, IEAUN, Portici; 1 9, BMNH, London); Mysore,
on coconut roots (Palmae), 21.vi.1937 (7. V. Subramaniam) (3 first stage, BMNH, London).
Ma taysIA: Malaya, Penang, 1 9 (IEAUN, Portici). VIETNAM: 1 9, Than-hoa (IEAUN, Portici).
HonG Kona: | 9, Repulse Bay (IEAUN, Portici).
Discussion. This is an unusual species that will probably be found underground throughout
south-east Asia, attended by the ant Acropyga acutiventris. The peculiar antennae, with the unusual
articulation between the first and second segments, are similar in all stages but as yet no second
stage has been available.
References
Balachowsky, A. S. 1957. Sur un nouveau genre aberrant de cochenille radicicole myrmécophile nuisible
au caféier en Colombie. Revue Path. vég. Ent. agric. Fr. 36: 157-164, 1 fig.
Beardsley, J. W. 1970. Three new species of Chavesia Balachowsky from Tropical America (Homoptera :
Coccoidea). Proc. Hawaii. ent. Soc. 20: 509-520, 5 figs.
Bowers, W. S., Nault, L. R., Webb, R. E. & Dutky, S. R. 1972. Aphid alarm pheromone: isolation, identi-
fication, synthesis. Science, N.Y. 177: 1121-1122.
Broadbent, L. 1951. Aphid excretion. Proc. R. ent. Soc. Lond. (A) 26: 97-103.
Biinzli, G. H. 1935. Untersuchungen iiber coccidophile Ameisen aus den Kaffeefeldern von Surinam.
Mitt. schweiz. ent. Ges. 16: 453-593, 51 figs.
Callow, R. K., Greenway, A. R. & Griffiths, D. C. 1973. Chemistry of the secretion from the cornicles of
various species of aphids. J. Insect Physiol. 19 : 737-748, 3 figs.
Chen, Su-Wan & Edwards, J. S. 1972. Observations on the structure of secretory cells associated with
aphid cornicles. Z. Zellforsch. mikrosk. Anat. 130: 312-317, 3 figs.
Dahl, M. L. 1971. Uber einen Schreckstoff bei Aphiden. Dr. ent. Z. (N.F.) 18: 121-128.
Das, G. M. 1959. Observations on the association of ants with Coccids of tea. Bull. ent. Res. 50 : 437-448,
6 figs.
Delage-Darchen, B., Matile-Ferrero, D. & Balachowsky, A. S. 1972. Sur un cas aberrant de symbiose.
Cochenilles x Fourmis. C. r. hebd. Séanc. Acad. Sci., Paris (D) 275 : 2359-2361, 1 pl.
De Lotto, G. 1977. On some African mealybugs (Homoptera: Coccoidea: Pseudococcidae). J. ent. Soc.
sth. Afr. 40 : 13-36, 6 figs.
Dixon, A. F. G. 1958. The protective function of the siphunculi of the nettle aphid, Microlophium evansi
(Theob.) (Hem., Aphididae). Entomologist’s mon. Mag. 94: 8.
—— & Stewart, W. A. 1975. Function of the siphunculi in aphids with particular reference to the syca-
more aphid, Drepanosiphum platanoides. J. Zool., Lond. 175 : 279-289, 4 figs.
Edwards, J. S. 1966. Defense by smear: supercooling in the cornicle wax of aphids. Nature, Lond. 211:
73-74.
Ferris, G. F. & Murdock, G. E. 1936. Contributions to the knowledge of the Coccoidea (Homoptera).
Ill. Microentomology 1: 115-122, 3 figs.
Flanders, S. E. 1957. The complete interdependence of an ant and a Coccid. Ecology 38 : 535-536.
Greenway, A. R. & Griffiths, D. C. 1973. A comparison of triglycerides from aphids and their cornicle
secretions. J. Insect Physiol. 19 : 1649-1655, 2 figs.
Hottes, F. C. 1928. Concerning the structure, function, and origin of the cornicles of the family Aphididae.
Proc. biol. Soc. Wash. 41 : 71-84, 30 figs.
Kislow, C. J. & Edwards, L. J. 1972. Repellent odour in aphids. Nature, Lond. 235 : 108-109.
Koteja, J. 1974a. Comparative studies on the labium in the Coccinea (Homoptera). Zesz. nauk. Akad.
roln. Krakow 27: 1-162, 78 figs.
1974b. The occurrence of a campaniform sensillum on the tarsus in the Coccinea (Homoptera).
Polskie Pismo ent. 44: 243-252, 3 figs.
—— 1974c. On the phylogeny and classification of the scale insects (Homoptera, Coccinea) (discussion
based on the morphology of the mouthparts). Acta zool. cracov. 19 : 267-325, 5 figs.
70
Le Pelley, R. H. 1943. An oriental mealybug (Pseudococcus lilacinus Ck\l.) and its insect enemies. Trans.
R. ent. Soc. Lond. 93 : 73-93, 7 figs, 4 pls.
Lindsay, K. L. 1969. Cornicles of the pea aphid, Acyrtosiphon pisum; their structure and function. A light
and electron microscope study. Ann. ent. Soc. Am. 62: 1015-1021, 6 figs.
Lloyd D. C. & Martini, E. 1957. A note on the circulus as an adhesive organ in some Pseudococcidae.
Can. Ent. 89: 46-48.
Nault, L. R. & Bowers, W. S. 1974. Multiple alarm pheromones in aphids. Entomologia exp. appl. 17:
455-457.
—— Edwards, L. J. & Styer, W. E. 1973. Aphid alarm pheromones: secretion and reception. Envir. Ent.
2: 101-105, 3 figs.
Newstead, R. 1903. Monograph of the Coccidae of the British Isles. 2. viii+ 270 pp., 4 figs, 52 pls. London.
Nixon, G. 1951. The association of ants with Aphids and Coccids. 36 pp. London.
Pesson, P. 1939a. Une glande exsertile ventrale chez les Coccides. Bull. Soc. zool. Fr. 64: 144-152, 4 figs.
—— 1939b. Ostioles et autohémorrhée chez les Coccides. Bull. Soc. zool. Fr. 64: 152-159, 2 figs.
Prins, A. J., Ben-Dov, Y. & Rust, D. J. 1975. A new observation on the association between ants
(Hymenoptera : Formicidae) and armoured scale insects (Homoptera : Diaspididae). J. ent. Soc. sth.
Afr. 38 : 211-216, 6 figs.
Reyne, A. 1954. Hippeococcus a new genus of Pseudococcidae from Java with peculiar habits. Zool.
Meded. Leiden 32 : 233-257, 53 figs.
Silvestri, F. 1924. A new myrmecophilous genus of Coccidae (Hemiptera) from India. Rec. Indian Mus.
26 : 311-315, 5 figs.
—— 1926. Descrizione di un novo genere di Coccidae (Hemiptera) mirmecofilo della Cina. Boll. Lab.
Zool. gen. agr. R. Scuola Agric. Portici 18 (1925) : 271-275, 2 figs.
— 1927. Descriptions of two species of myrmecophilous Coccidae (Hemiptera Insecta). China J.
7: 253-254, 2 figs.
Smith, M. R. 1942. The relationship of ants and other organisms to certain scale insects on coffee in Puerto
Rico. J. Agric. Univ. P. Rico 26: 21-27.
Strickland, A. H. 1951. The entomology of Swollen Shoot of cacao. I — the insect species involved, with
notes on their biology. Bull. ent. Res. 41 : 725-748, 1 fig.
Strong, F. E. 1969. Observations on aphid cornicle secretions. Ann. ent. Soc. Am. 60: 668-673, 4 figs.
Sulc, K. 1909. Zur Anatomie der Cocciden. Zool. Anz. 34: 164-172, 4 figs.
Takahashi, R. 1934. Observations on the Coccidae of Formosa. Part IV. Rep. Govt res. Inst. Dep. Agric.
Formosa 63: 1-38, 24 figs.
— 1941. Two new myrmecophilous Coccidae from Thailand (Siam). Tenthredo 3 : 201-207, 2 figs.
—— 1942. Some injurious insects of agricultural plants and forest trees in Thailand and Indo-China,
II. Coccide. Rep. Govt Res. Inst. Dep. Agric. Formosa 81 : 1-56, 26 figs.
—— 1950. Some species of Coccidae from the Riouw Islands. Part I. Insecta matsum. 17 : 65-72, 5 figs.
— 1951. Three new myrmecophilous scale insects from Malay Peninsular (Homoptera). Mushi 22:
1-8, 4 figs.
Way, M. J. 1954. Studies on the association of the ant Oecophylla longinoda (Latr.) (Formicidae) with the
scale insect Saissetia zanzibarensis Williams (Coccidae). Bull. ent. Res. 45: 113-134, 8 figs.
—— 1963. Mutualism between ants and honeydew-producing Homoptera. A. Rev. Ent. 8 : 307-344.
Wheeler, W. M. 1921. A study of some social beetles in British Guiana and of their relations to the ant-
plant Tachigalia. Zoologica, N.Y. 3: 35-126, 12 figs, 5 pls.
Williams, D. J. 1969. The family-group names of the scale insects (Hemiptera : Coccoidea). Bull. Br. Mus.
nat. Hist. (Ent.) 23 : 315-341.
—— 1970. The mealybugs (Homoptera, Coccoidea, Pseudococcidae) of sugar-cane, rice and sorghum.
Bull. ent. Res. 60: 109-188, 29 figs.
Wilson, E. O. 1971. The insect societies. xii+548 pp. Cambridge, Massachusetts.
Wynn, G. G. & Boudreaux, H. B. 1972. Structure and function of aphid cornicles. Ann. ent. Soc. Am.
65 : 157-166, 13 figs.
Index
acariformis, 15 brevipes, 10 Dysmicoccus, 10
Allomyrmococcini, 14
Allomyrmococcus, 14, 15 Chavesia, 6, 7, 13, 62 Ehrhornia, 9
annandalei, 66 chiengraiensis, 56 Eumyrmococcus, 62, 63
Antonina, 9, 10 Chnaurococcus, 12 Euripersia, 12
71
europaea, 12
farinosus, 10
formicarii, 31, 33
Hippeococcus, 14, 15, 20, 22
khooi, 31, 39
lilacinus (Planococcus), 8
lilacinus (Pseudococcus), 8
longispinus, 10
Malaicoccus, 14, 15, 30, 31
montanus, 22
moundi, 31, 44
obscurus, 7
Paramyrmococcus, 14, 15, 54
Phenacoccus, 10
Pseudococcinae, 9, 14
Pseudococcus, 7, 8, 10
rappardi, 22
Rhizoecinae, 9, 62
Rhodania, 9
riouwensis, 33, 50
fe
scorpioides, 66
smithii, 63
Sphaerococcinae, 9
subterraneus, 12
takahashii, 31, 52
Trabutininae, 9
trinidadensis, 13
vietnamensis, 56, 61
wegneri, 22, 26
Xenococcus, 62, 63
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% GENERAL 4
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Bulletin of the /
British Museum (Natural History)
A supplementary catalogue of the family-
sroup and genus-group names of the
Coleophoridae (Lepidoptera)
K. Sattler & W. G. Tremewan
Entomology series Vol37No2 25 May 1978
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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© Trustees of the British Museum (Natural History), 1978
ISSN 0524-6431 Entomology series
Vol 37 No 2 pp 73-96
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 May 1978
genus-group names of the Coleophoridae
(Lepidoptera)
K. Sattler & W. G. Tremewan
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis , ; : ; : ; : : . ; : ; , 73
Introduction . ‘ : ; , ; ‘ : : F ‘ : : 7B
Acknowledgements . ; : : * : : : . : ; : 74
The classification of the Coleophoridae by Capuse and Falkovitsh d ; s 4
Check-list of Coleophoridae following Capuse (1973; 1975) . ; : ‘ reamney
The synonymies of Coleophora sternipennella (Zetterstedt, [1839]) and C. benanderi
Kanerva, 1941 : ; : P ; : . : : : : Ss - ST
The identity of Phalaena (Tinea) vestianella Linnaeus, 1758 . , : ‘ . 84
Alphabetical catalogue of the family-group names : : : . ‘ gt
Alphabetical catalogue of the genus-group names ’ ; : : ; . 86
References : ; : . ' : : : ' : : * 3. 92
Index. : ‘ : : 5 : : . : : é , : 93
Synopsis
The family-group and genus-group names (including variations in spelling) of the Coleophoridae pub-
lished since 1973 are listed alphabetically. Citations of type-species are given, together with biblio-
graphical references to the original descriptions and designations of type-species. The recent publications
of Capuse and Falkovitsh on the classification of the Coleophoridae are discussed. One subtribal, thirty-
seven generic and three specific synonymies are newly introduced. The identity of Phalaena (Tinea)
vestianella Linnaeus is established and a lectotype is designated. The synonymies of three confused
species are discussed and the male genitalia are illustrated.
Introduction
Recently, Capuse (1971) and Falkovitsh (1972) proposed a revised classification of the Coleo-
phoridae. The main feature of this classification, which fundamentally differs little from the earlier
system (Toll, 1953; 1962), is the application of numerous generic and suprageneric names to what
were previously considered to be merely species-groups. In 1974 we discussed this classification
and catalogued the family-group and genus-group names of the Coleophoridae. While our
catalogue was in press, and after its publication, further genus-group names were proposed
(Capuse, 1973; 1974; 1975) and, because of the large number of new taxa, we find it necessary
to review these subsequent papers. Our earlier criticism can be equally applied to these new taxa
and our opinion on the validity of the divisions of the Coleophoridae, which we expressed in
detail (Sattler & Tremewan, 1974 : 186-191), remains unchanged.
Each generic name has been checked for homonymy in the catalogues of Neave (1939-66,
Nomencl. zool. 1-6). Names that have been proposed expressly to replace junior homonyms are
referred to in this catalogue as objective replacement names. Subjective synonymy of the genera
is not discussed in detail; however, the present status of each genus is recorded. oe
All references have been checked personally by the authors. In all instances the original
Bull. Br. Mus. nat. Hist. (Ent.) 37 (2): 73-96 Issued 25 May 1978
73
publications were examined because reprints sometimes differ in date of publication and in
pagination. The printed date of publication in a book or journal is accepted as correct, unless
there exists published evidence to the contrary.
The format of this supplement follows that of our previous catalogue. Family-group and
genus-group names are listed in separate sections. All names are arranged in alphabetical order;
synonyms and unavailable names are cross-referenced. Junior objective synonyms and unavail-
able names (nomina nuda) are in non-bold italics; unavailable names are marked with a double
dagger ({). The alphabetical entries of all other generic names are in bold italics, as are the
names of their type-species.
Acknowledgements
We gratefully acknowledge the help of Mr R. G. Fennah, Commonwealth Institute of Entomology,
London; Mr H. Patzak, Aschersleben; Mr B. W. Rasmussen, Universitetets Zoologiske Museum,
Copenhagen; Dr G. S. Robinson and Mr P. E. S. Whalley, British Museum (Natural History),
London.
The classification of the Coleophoridae by Capuse and Falkovitsh
In his major paper on the Coleophoridae, Capuse (1971) divided this family into numerous sub-
families, tribes, subtribes, genera and subgenera without providing any keys. However, keys
were subsequently provided in a privately published paper (Capuse, 1973), in which a sub-
tribe, 28 genera and 8 subgenera were erected. In this paper the short introduction (p. 3) is fol-
lowed by a key to the subfamilies of the Coleophoridae (pp. 4-5) and a key to the tribes of the
subfamily Coleophorinae (pp. 5-6), the other subfamilies being monobasic. Three separate keys
to the genera of the tribes Carpochenini (pp. 7-8), Casignetini (pp. 8-12) and Coleophorini
(pp. 12-21) are provided. A further key is entitled ‘Clé de determination des genres dont nous
n’avons pas examiné l’espéce type’ (Key for the determination of the genera of which we have
not examined the type-species) (pp. 21-23) and is followed by the bibliography (p. 24).
The new genera and subgenera proposed in the keys are accompanied by type-species desig-
nations and satisfy the provisions of the International Code of Zoological Nomenclature. How-
ever, it would have been desirable for them to have been accompanied by detailed descriptions,
illustrations, a discussion of the related taxa, an indication of their exact position in Capuse’s
system and a list of further included species. With the exception of one genus all the new genera
and subgenera are monotypic, which seems to indicate that they were based either on the study
of a small number of species or on a very narrow concept of the genus.
Although Capuse placed the new genera and subgenera in their appropriate tribes he did not
discuss their affinities with other genera — that two genera key out next to each other is not
necessarily an indication of close relationship.
The taxa named in the keys were further described in a subsequent paper (Capuse, 1975)
where they are again marked as ‘nov.’. However, it is wrong and misleading to indicate them as
new because they were made nomenclaturally available in 1973. In addition, Capuse made no
reference to his earlier publication except for a brief mention in the introduction and a citation
in the bibliography, and nowhere does he give an indication that any of the taxa were newly
proposed in the keys of 1973.
We have already discussed in detail the characters used by Capuse and Falkovitsh for separating
taxa. These characters were found to be unacceptable at the generic and suprageneric level and,
frequently, are merely species differences; their use resulted in poorly defined genera with transi-
tions between each other. Capuse’s work is based mainly on the papers of Toll (1953; 1962) and
this he should have clearly stated. In many instances the groups, sections and subsections of Toll
were fully recognized by Capuse who merely gave them names and raised them to generic or even
suprageneric rank, sometimes rearranging the species-groups and slightly altering the sequence
of the species within the groups. The mere application of names to widely recognized species-
groups contributes nothing to the advancement of science.
74
The genus Cornulivalvulia Capuse, 1973, originally described in the tribe Casignetini Falko-
vitsh, was subsequently synonymized by Falkovitsh (1974a: 194; 1974b : 138) with the genus
Multicoloria Capuse, 1973, originally described in the tribe Coleophorini Hiibner. This synonymy
further emphasizes the weakness of the system proposed by Capuse and Falkovitsh.
The division of the Coleophoridae and particularly the genus Coleophora into subfamilies,
_ tribes, subtribes, genera and subgenera by Capuse (1971) and Falkovitsh (1972) has been ignored
by all subsequent authors (e.g. Patzak (1974) in his important paper on the casebearers of East
Germany) with the exception of Reznik (1976), who recognized the genus Multicoloria Capuse.
However, it is apparent from Reznik’s bibliography that he was unaware of our critical treatment
of the Coleophoridae (Sattler & Tremewan, 1974). After the division of the Coleophoridae by
Capuse (1971) and Falkovitsh (1972), the latter (Falkovitsh, 1973), in a major paper on the case-
bearers of the Kysyl Kum, in which he described numerous new species, used the name Coleo-
phora throughout; however, his paper may have been delayed in press.
As the original works of Capuse and Falkovitsh were only based on about 130 of an estimated
total of 1000 species, it seemed inevitable to us that the study of further species would result in
the erection of more new genera. If the narrow genus-concept of Capuse and Falkovitsh is not
rejected, the separation of further large numbers of genera (200 or more!) from Coleophora is
unavoidable. We do not believe that such drastic splitting would serve any useful purpose and
suggest that no further attempt is made to divide Coleophora.
Check-list of Coleophoridae following Capuse (1973; 1975)
The following check-list contains the names of the taxa dealt with by Capuse (1973; 1975); the
arrangement follows his 1975 paper. The groups, sections and subsections of Toll (1953; 1962)
are also indicated. Obvious misspellings and wrong authors have been corrected. The names
atticae, herzegoviniensis, rectivittella and subsimilis, all attributed to Toll by Capuse (1975),
originated from Toll but to our knowledge were never made nomenclaturally available. They
are not recorded in the British Museum (Natural History) card index or in a manuscript list of
the species described by Toll which was compiled by his wife after his death. These names are
nomina nuda and are marked with a double dagger ({). The name C. subechinella is used by Toll
(1962 : pl. 17S, fig. 183); however, it is not ‘accompanied by a statement that purports to give
characters differentiating the taxon’ (Jnt. Code zool. Nom., Article 13(a)(i)) and is therefore
unavailable.
COLEOPHORIDAE Hiibner, 1825
COLEOPHORINAE Hiibner, 1825
CASINI Capuse, 1971
Casas Wallengren, 1881
C. leucapennella (Hiibner) group 1 (1962)
COLEOPHORINI Hiibner, 1825
HAPLOPTILIINA Barnes & McDunnough, 1917
Postvinculia Capuse, 1973
P. lutipennella (Zeller) group 2, section 2 (1962)
Paravalvulia Capuse, 1973
P. spiraeella (Rebel) group 2, section 2 (1962)
Haploptilia Hiibner, 1825
H. coracipennella (Hiibner)
Zagulajevia Capuse, 1971
Z. tadzhikiella (Danilevski) group 7, section 2, subsection 2 (1962)
Frederickoenigia Capuse, 1971
F. flavipennella (Herrich-Schaffer) group 2, section 5 (1962)
Orghidania Capuse, 1971
O. gryphipennella (Bouché) group 2, section 4 (1962)
Sacculia Capuse, 1973
S. excellens (Toll) group 6, section 3 (1962)
S. subexcellens (Toll) group 6, section 3 (1962)
75
Characia Falkovitsh, 1972
C. haloxyli (Falkovitsh)
COLEOPHORINA Hiibner, 1825
Dumitrescumia Capuse, 1973
D. cecidophorella (Oudejans)
D. trigeminella (Fuchs)
D. hydrolapathella (Hering)
Tuberculia Capuse, 1973
T. albitarsella (Zeller)
Globulia Capuse, 1973
G. cornuta (Heinemann & Wocke)
Quadratia Capuse, 1973
Q. fuscocuprella (Herrich-Schaffer)
Tollsia Capuse, 1973
T. hornigi (Toll)
Kasyfia Capuse, 1973
K. binderella (Kollar)
K. orbitella (Zeller)
Rhamnia Capuse, 1973
R. ahenella (Heinemann)
R. subahenella Capuse
Oudejansia Capuse, 1973
. obviella (Rebel)
. moestella (Toll)
. pseudoobviella (Toll)
. Striolatella (Toll)
. cribrella (Toll)
. dichroella (Toll)
. cuencella (Toll)
. obtectella (Zeller)
. paraobviella (Toll)
Membrania Capuse, 1973
M. (M.) calycotomella (Stainton)
M. (M.) trifisella (Rebel)
Longibacillia Capuse, 1973
M. (L.) fergana (Toll)
M. (L.) schahkuhensis (Toll)
M. (L.) niveopictella (Toll)
Baraschia Capuse, 1973
B. (B.) paradoxella (Toll)
B. (B.) uralensis (Toll)
Abaraschia Capuse, 1973
B. (A.) pagmana (Toll)
Kutznetzovvlia Capuse, 1973
K. solidaginella (Staudinger)
K. conyzae (Zeller)
K. angustilinea (Toll)
K. mellechella (Toll)
K. semistriatella (Toll)
K. ptarmica (Walsingham)
K. namangana (Toll)
K. santolinella (Constant)
K. jerusalemella (Toll)
K. parcella (Toll)
Glaseria Capuse, 1971
G. (G.) biseriatella (Staudinger)
G. (G.) microxantha (Walsingham)
Proglaseria Capuse, 1973
G. (P.) laticostella (Mann)
OSOSO9 9009
76
group 2, section 8 (1962)
group 2, section 8 (1962)
group 2, section 8 (1962)
group 3 (1962)
group 2, section 11, subsection 1 (1962)
group 2, section 11, subsection 2 (1962)
group 2, section 14 (1962)
group 2, section 15 (1962)
group 2, section 15 (1962)
group 2, section 15 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 2 (1962)
group 6, section 4, subsection 1 (1962)
group 6, section 4, subsection 1 (1962)
group 6, section 4, subsection 2 (1962)
group 6, section 4, subsection 2 (1962)
group 6, section 4, subsection 2 (1962)
group 6, section 5, subsection 1 (1962)
group 6, section 5, subsection 1 (1962)
group 6, section 5, subsection 2 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group. 6, section 1 (1962)
group 6, section 1 (1962)
group 6, section 1 (1962)
group 9, section 3 (1962)
group 9, section 3 (1962)
group 9, section 2 (1962)
G. (P.) murciana (Toll)
G. (P.) fretella (Zeller)
G. (P.) pabulella (Zeller)
G. (P.) marianii (Toll)
G. (P.) perplexella (Toll)
G. (P.) gilveolella (Toll)
G. (P.) xanthoptera (Toll)
Scleriductia Capuse, 1973
S. ochripennella (Zeller)
Ascleriductia Capuse, 1973
A. lithargyrinella (Zeller)
A. fuscatella (Toll)
Helvalbia Capuse, 1973
H. lineolea (Haworth)
Latisacculia Capuse, 1973
L. crocinella (Tengstr6m)
L. arenariella (Zeller)
L. medelichensis (Krone)
L. protecta (Walsingham)
L. pseudoserenella (Toll)
Klimeschja Capuse, 1973
(Klimeschjosefia Capuse, 1975)
K. oriolella (Zeller)
K. vulnerariae (Zeller)
K. semicinerea (Staudinger)
K. medicaginella (Mann)
K. defessella (Herrich-Schaffer)
K. mongetella (Chrétien)
K. siliquella (Constant)
K. coarctella (Staudinger)
K. femorella (Walsingham)
K. albicosta (Haworth)
K. supinella (Ortner)
K. suboriolella (Toll)
K. hartigi (Toll)
K. bruneosignata (Toll)
K. rudella (Toll)
K. fraudulentella (Toll)
K. rugulosa (Toll)
K. turbatella (Toll)
K. glaseri (Toll)
_ Amseliphora Capuse, 1971
A. (A.) niveicostella (Zeller)
A. (A.) albicostella (Duponchel)
A. (A.) squamella (Constant)
A. (A.) trifariella (Zeller)
A. (A.) bilineella (Herrich-Schiaffer)
. (A.) saturatella (Stainton)
. (A.) acrisella (Milliére)
. (A.) imbecilla (Toll)
. (A.) discordella (Zeller)
. (A.) genistae (Stainton)
A. (A.) onobrychiella (Zeller)
A. (A.) rectilineella (Fischer von
Roeslerstamm)
A. (A.) albidorsella (Toll)
A. (A.) lineata (Toll)
A. (A.) algeriensis (Toll)
A. (A.) paragenistae (Kasy)
A
A
A
A
A
group 9, section 2 (1962)
group 9, section 2 (1962)
group 9, section 2 (1962)
group 9, section 2 (1962)
group 9, section 2 (1962)
group 9, section 2 (1962)
group 9, section 2 (1962)
group 2, section 3 (1962)
group 9, section 1 (1962)
group 9, section 1 (1962)
group 6, section 8 (1962)
group 9, section 1 (1953)
group 9, section 1 (1953)
group 23 (1953)
group 23 (1953)
group 9, section 3, subsection B (1953)
group 9, section 3, subsection C (1953)
group 9, section 2 (1953)
group 9, section 3, subsection B (1953)
group 9, section 2 (1953)
group 9, section 3, subsection A (1953)
Ardania Capuse, 1973
A. (A.) bilineatella (Zeller) group 9, section 2 (1953)
Amselghia Capuse, 1973
A. (A.) fringillella (Zeller) group 9, section 3, subsection A (1953)
Calcomarginia Capuse, 1973
C. ballotella (Fischer von Roeslerstamm) group 14 (1953)
Apista Hiibner, 1825
A. iranella (Toll)
A. escalerai (Toll)
A. albostraminata (Toll)
A. arenicola (Toll)
A. ochristrigella (Ragonot)
Apista Hiibner, 1825, s. str.
A. (A.) gallipennella (Hiibner) group 18, section 1, subsection A (1953)
Metapista Capuse, 1973
A. (M.) stramentella (Zeller) group 18, section 1, subsection B (1953)
A. (M.) colutella (Fabricius)
A. (M.) rebeli (Gerasimov)
A. (M.) wiltshirei (Toll)
A. (M.) agnatella (Toll)
A. (M.) kasyi (Toll)
Multicoloria Capuse, 1973
M. ditella (Zeller) group 18, section 4 (1953)
M. conspicuella (Zeller) group 18, section 3 (1953)
M. tristraminata (Toll)
M. similis (Staudinger)
M. vibicigerella (Zeller) group 18, section 4 (1953)
M. partitella (Zeller) group 18, section 4 (1953)
M. pirizanella (Toll)
M. andalusiae (Toll)
M. leucostrigella (Toll)
M. satellitella (Toll)
M. medicaginis (Herrich-Schaffer) group 18, section 4 (1953)
M. zonatella (Toll)
M. anatolica (Toll)
M. eucera (Toll)
M. astragalella (Zeller) group 18, section 5 (1953)
M. caelebipennella (Zeller) group 18, section 7 (1953)
M. mandschuriae (Toll)
M. polonicella (Zeller) group 18, section 7 (1953)
M. vibicella (Hiibner) group 18, section 7 (1953)
M. didyma (Toll)
M. stachi (Toll)
M. paraspumosella (Toll)
M. spumosella (Toll)
M. dubiosa (Toll)
M. xanthoargentea (Toll)
M. involucrella (Chrétien)
M. qulikushella (Toll)
M. paraononidella (Amsel)
M. ononidella (Milliére)
M. giraudi (Constant)
M. paragiraudi (Toll)
M. depauperella (Toll)
M. perserenella (Rebel)
M. semistrigata (Toll)
M. agrianella (Rebel)
M. canariipennella (Toll)
M. simillima (Staudinger)
78
M.
M.
M.
M.
fuscociliella (Zeller)
menephilella (Toll)
helichrysella (Krone)
biskraensis (Toll)
M. radiosella (Toll)
M.
M.
M.
M.
M.
M.
M.
M.
M.
dubiella (Baker)
minaxella (Toll)
praeposita (Toll)
atlanticolella (Zerny)
argyrella (Herrich-Schaffer)
flabelligerella (Rebel)
amasiella (Staudinger)
sumptuosa (Toll)
honestella (Toll)
M. gazella (Toll)
M. gozmanyi (Toll)
M.
M.
M.
M.
M.
M.
. valesianella (Zeller)
. echinella (Staudinger)
. berlandella (Toll)
. tozeurensis (Toll)
. tsubsimilis Capuse, nomen nudum
. trectivittella Capuse, nomen nudum
. tsubechinella (Toll)
. cartilaginella (Christoph)
SESSSESS
albarracinica (Toll)
medicagivora (Toll)
cyrniella (Rebel)
praecipua (Walsingham)
fuscostraminella (Toll)
microeucera (Toll)
Damophila Curtis, 1832
be
D.
D.
Dp:
D.
Ds
D. therzegoviniensis Capuse, nomen nudum
Dy
Dp;
spissicornis (Haworth)
hieronella (Zeller)
fuscicornis (Zeller)
fuscoaenea (Toll)
amethystinella (Ragonot)
leucostoma (Gerasimov)
tatticae Capuse, nomen nudum
dannehli (Toll)
Lucidaesia Capuse, 1973
L. frischella (Linnaeus)
Pe eb
. trifolii (Curtis)
. cuprariella (Zeller)
. cuprifulgella (Toll)
. auronitella (Toll)
. argentifimbriata (Walsingham)
. basimaculella (Mann)
CASIGNETINI Falkovitsh, 1972
CASIGNETINA Falkovitsh, 1972
Cornulivalvulia Capuse, 1973
C.
vicinella (Zeller)
Aureliania Capuse, 1971
AAA A RAD
(A.) flavaginella (Zeller)
(A.) virgaureae (Stainton)
(A.) halophylella (Zimmermann)
(A.) therinella (Tengstrém)
(A.) sternipennella (Zetterstedt)
(A.) versurella (Zeller)
79
group 13, section 2 (1953)
group 5, section 1 (1962)
group 5, section 1 (1962)
group 5, section 1 (1962)
group 5, section 1 (1962)
group 5, section 1 (1962)
group 5, section 1 (1962)
group 5, section 2 (1962)
group 18, section 6 (1953)
group 30, section 2, subsection E (1953)
group 30, section 2, subsection A (1953)
group 30, section 2, subsection B (1953)
group 30, section 2, subsection C (1953)
group 30, section 2, subsection E (1953)
group 30, section 2, subsection E (1953)
. (A.) pratella (Zeller) group 30, section 2, subsection C (1953)
. (A.) motacillella (Zeller) group 30, section 2, subsection E (1953)
. (A.) erigerella (Ford)* group 30, section 2, subsection E (1953)
. (A.) leucogrammella (Herrich-Schaffer)
. (A.) luteolella (Staudinger)
. (A.) dentiferella (Toll)
. (A.) palumbipennella (Toll)
. (A.) mendosella (Toll)
. (A.) hsiaolingella (Toll)
. (A.) fiorii (Toll)
. (A.) lewandowskii (Toll) group 30, section 2, subsection A (1953)
. (A.) cinerea (Toll)
Nosyrislia Capuse, 1973
A. (N.) linosyris (Hering) group 30, section 2, subsection D (1953)
A. (N.) asteris (Miihlig) group 30, section 2, subsection D (1953)
Ecebalia Capuse, 1973
A. (E.) laripennella (Zetterstedt) group 30, section 2, subsection F (1953)
A. (E.) parenthella (Toll)
A. (E.) gaviaepennella (Toll)
A. (E.) botauripennella (Toll)
Hamuliella Capuse, 1973
AA AAAAARAAAA
H. otitae (Zeller) group 30, section 4, subsection E (1953)
H. granulatella (Zeller sensu Klimesch) group 30, section 4, subsection C (1953)
H. galatellae (Hering) group 30, section 4, subsection F (1953)
H. klemensiewiczi (Toll) group 30, section 4, subsection D (1953)
H. dianthivora (Walsingham)
H. scabrida (Toll)
H. agenjoi (Toll)
H. griseomixta (Toll)
H. heringi (Toll)
H. obliterata (Toll)
H. opacella (Toll)
H. insulicola (Toll)
H. gnaphalii (Zeller) group 30, section 4, subsection C (1953)
H. fischeri (Toll)
H. tolli (Klimesch)
H. succursella (Herrich-Schaffer) group 30, section 4, subsection B (1953)
H. separatella (Benander) group 30, section 4, subsection A (1953)
H. riffelensis (Rebel)
H. sabulella (Toll)
H. granulosella (Staudinger)
H. artemisicolella (Bruand) group 30, section 3, subsection D (1953)
Neugenvia Capuse, 1973
N. vlachi (Toll) group 30, section 6, subsection A (1953)
N. peribenanderi (Toll) group 30, section 4, subsection G (1953)
N. lineariella (Zeller) group 30, section 5 (1953)
N. ramosella (Zeller)
N. troglodytella (Duponchel) group 30, section 6, subsection B (1953)
N. axana (Hering)
N. asterifoliella (Klimesch)
N. gardesanella (Toll)
Benanderpia Capuse, 1973
B. adspersella (Benander) group 30, section 10, subsection B (1953)
B. fulvosquamella (Herrich-Schaffer) group 30, section 10, subsection A (1953)
Patzakia Capuse, 1973
P. silenella (Herrich-Schaffer) group 30, section 10, subsection D (1953)
P. robustella (Fuchs)
pe piece Ford, 1935, is currently considered to be a synonym of C. squamosella Stainton, 1856 (Bradley,
: 181).
80
P. ciconiella (Herrich-Schiaffer)
P. tritici (Lindemann)
P. graminicolella (Heinemann)
Lvaria Capuse, 1973
L. lassella (Staudinger) group 30, section 11 (1953)
Ulna Capuse, 1973
U. saponariella (Heeger) group 30, section 13 (1953)
GONIODOMINA Capuse, 1973
Goniodoma Zeller, 1849
G. auroguttella Zeller
Klinzigedia Capuse, 1971
K. phlomidella (Christoph)
CARPOCHENINI Capuse, 1973
Carpochena Falkovitsh, 1972
C. squalorella (Zeller) group 34 (1953)
Ionescumia Capuse, 1971
I. clypeiferella (Hofmann) group 34 (1953)
Stollia Capuse, 1971
S. binotapennella (Duponchel) group 34 (1953)
Bourgogneja Capuse, 1971
B. onosmella (Brahm) group 29 (1953)
The synonymies of Coleophora sternipennella (Zetterstedt, [1839])
and C. benanderi Kanerva, 1941
The misidentified type-species of Aureliania Capuse, 1971, viz. Coleophora flavaginella Lienig &
Zeller sensu Capuse, and the type-species of Bacescuia Capuse, 1971, viz. moeniacella Stainton,
are both involved in complicated and confusing synonymy which we attempt to clarify below. The
species Phalaena (Tinea) vestianella Linnaeus is also involved (see pp. 84-85). To avoid further
confusion we illustrate the ¢ genitalia of the three species concerned. In the synonymies reference
is also made to genitalia figures in important papers on Coleophoridae.
Coleophora sternipennella (Zetterstedt, [1839])
(Fig. 1)
Ornix sternipennella Zetterstedt, [1839], Insecta lapponica : 1011. Syntypes 1 3, 1 29, SWEDEN: Vasterbotten,
Lycksele, 23.vi.1832 (Zetterstedt) (Universitetets Zoologiska Institution, Lund) [not examined].
Coleophora flavaginella Lienig & Zeller, 1846, Isis, Leipzig 1846:295. LECTOTYPE 4, U.S.S.R.:
Latvia (‘Lievland’), Kokenhusen near Riga (Lienig) (genitalia slide no. 8381; British Museum (Natural
History), London), here designated [examined].
Coleophora {flavagipennella Lienig & Zeller, 1846, Isis, Leipzig 1846: 295. [A manuscript name of
Fischer von Roeslerstamm; first published in synonymy under flavaginella Lienig & Zeller and there-
fore unavailable (Int. Code zool. Nom., Article 11(d)).]
Coleophora punctipennella Nylander, 1848, in Tengstrém, Notis. Sdllsk. Faun. Fl. fenn. Forh. 1: 142.
Syntype(s), FINLAND: Helsinki (‘Helsingfors’), vii. (Nylander) (Zoological Museum, Helsinki) [not
examined].
Coleophora muehligiella Stainton, 1887, Entomologist’s mon. Mag. 24: 14. LECTOTYPE 9, [? GERMANY:
Frankfurt/Main] (Miihlig) (genitalia slide no. 11093; British Museum (Natural History), London),
here designated [examined].
Coleophora moeniacella Stainton, 1887, Entomologist’s mon. Mag. 24: 42. [Objective replacement name
for C. muehligiella Stainton, 1887.]
a eye sternipennella (Zetterstedt); Benander, 1939, Opusc. ent. 4: 95, pl. 5, figs 73, 73a (3), pl. 6,
g. 85 (2).
Eupista sternipennella (Zetterstedt); Toll, 1953, Mater. Fizjogr. Kraju 32: 177, pl. 14, figs 122, 122a (),
pl. 29, fig. 278 (9).
Aureliania sternipennella (Zetterstedt) Capuse, 1971, Recherches morph. syst. Famille Coleophoridae : pl.
23, fig. E, pl. 38, fig. E.
81
Coleophora sternipennella (Zetterstedt); Bradley, 1972, Handbk Ident. Br. Insects 11 (2) : 16.
Coleophora sternipennella (Zetterstedt); Patzak, 1974, Beitr. Ent. 24: 264, colour fig. 32 (forewing),
figs 7 (case), 200 (3), 332 (9).
Coleophora sternipennella (Zetterstedt); Karsholt & Nielsen, 1976a, Syst. Fortegnelse over Danmarks
Sommerfugle : 28.
The syntypes of sternipennella were examined by Benander (1938 : 117; 1939 : 95, 99 [explanation
of pl. 6, fig. 85], pl. 6, fig. 85; 1940 : 62).
The specimen here designated as the lectotype of flavaginella originated from Lienig and came
into Stainton’s possession via Tengstrém and Nylander. It bears the label ‘Flavaginella Nylander
7/59’ indicating the date when Stainton received the specimen from Nylander (Stainton, 1887:
14). One 3, two 2 and one specimen without abdomen from the original series (coll. Zeller; coll.
Stainton; British Museum (Natural History), London) have been labelled as paralectotypes.
Pierce & Metcalfe (1938 : 67) stated that they had examined the genitalia of the ‘type’ of flava-
ginella. In the British Museum (Natural History) there is a specimen bearing several ‘Type’
labels and a label in Zeller’s handwriting ‘Flavaginella Lg. Is. 1846, 295.’. It lacks the abdomen
and, although it bears no slide number, is probably the specimen referred to by Pierce & Metcalfe;
Figs 1-3. Male genitalia of Coleophora species. (1) C. sternipennella (Zetterstedt), London, Bucking-
ham Palace Gardens, 26.vii.1974 (Bradley) (slide no. 20 402; BMNH). (2) C. benanderi Kanerva,
Dorset, Swanage, 8.vii.1897 (Bankes) (slide no. 4620; BMNH). (3) C. vestianella (Linnaeus),
[Sweden] (Linnaeus) (slide no. 715, Sattler; Linnean Society, London).
82
however, we have been unable to trace the genitalia slide preparation. This specimen is excluded
from the type-material as there is some doubt about its being authentic.
The syntypes of punctipennella were examined and the synonymy with sternipennella was
confirmed by Kanerva (1941 : 120).
In his original description of muehligiella, Stainton (1887 : 14) referred to specimens that he
had received from Miihlig in February, 1859; however, only a single female is in the Stainton
collection and there is no evidence in his notebook that he received more than one specimen.
Stainton proposed the name moeniacella ({maeniacella auctorum, incorrect subsequent spelling)
as an objective replacement name for Coleophora muehligiella Stainton, 1887, which he erroneously
considered to be preoccupied by Coleophora muehligella Wocke, [1876], in Heinemann, Schmett.
Dtl. Schweiz (2) 2 (2): 604. Therefore the names C. muehligiella Stainton and C. moeniacella
Stainton must always be placed together; in spite of this the two names have been separated by
subsequent authors.
When introducing the name atriplicis, Meyrick ([1928] : 763) stated ‘This species has been
erroneously identified with C. maeniacella Staint.’, at the same time placing ‘muehligiella Staint.,
non Wocke’ in synonymy of atriplicis. It should be noted that the name C. atriplicis must be
attributed to Meyrick, who made it nomenclaturally available, and not to Durrant from whom
it originated as a manuscript name.
Meyrick was followed by Pierce & Metcalfe (1935 : 64) who placed muehligiella Stainton in
synonymy under atriplicis but did not refer to moeniacella. In 1938, Pierce & Metcalfe (1938 : 67)
placed moeniacella [as ‘maeniacella’], together with annulatella, in synonymy under flavaginella
Lienig & Zeller.
Bradley (1972 : 16), in the British list, placed moeniacella [=muehligiella Stainton] as a senior
synonym of suaedivora Meyrick; however, Stainton (1887: 14, muehligiella) expressly dis-
tinguished it from the species on Suaeda maritima (L.) Dumortier (which he erroneously called
flavaginella Lienig).
Karsholt & Nielsen (1976a : 28) placed moeniacella as a synonym of sternipennella and muehli-
giella as a synonym of suaedivora Meyrick.
Coleophora benanderi Kanerva, 1941
(Fig. 2)
Coleophora benanderi Kanerva, 1941, Suomen hyént. Aikak. 7: 120. Syntypes 3, 2, SWEDEN: Schonen;
Blekinge; Smaland; Vastergétland; Oland; Gotland, vii-viii. (Benander) (coll. Benander, Sweden)
[not examined].
[Eupista annulatella (Nylander); Pierce & Metcalfe, 1935, Genitalia Tineid Families Lepid. Br. Islands : 65,
pl. 39, figs (g, 2). Misidentification.]
[Eupista flavaginella (Lienig & Zeller); Pierce & Metcalfe, 1938, Genitalia Br. Pyrales, Deltoids & Plumes :
67. Misidentification. ]
[Coleophora annulatella Nylander; Benander, 1939, Opusc. ent. 4: 96, pl. 5, figs 74, 74a (3), pl. 6, fig. 84
(2). Misidentification.]
[Eupista flavaginella (Lienig & Zeller); Toll, 1953, Mater. Fizjogr. Kraju 32: 177, pl. 14, figs 120, 120a
(3) (not pl. 29, fig. 277, 2). Misidentification.]
[Aureliania flavaginella (Lienig & Zeller) Capuse, 1971, Recherches morph. syst. Famille Coleophoridae :
65. Misidentification.]
[Coleophora annulatella Nylander; Bradley, 1972, Handbk Ident. Br. Insects 11 (2) : 16. Misidentification. ]
[Coleophora annulatella Nylander; Sattler & Tremewan, 1974, Bull. Br. Mus. nat. Hist. (Ent.) 30: 201.
Misidentification. ]
[Coleophora annulatella Nylander; Patzak, 1974, Beitr. Ent. 24: 263, figs 199 (3), 331 (9). Misidentifica-
tion.]
Coleophora benanderi Kanerva; Karsholt & Nielsen, 1976a, Syst. Fortegnelse over Danmarks Sommer-
Sugle : 28.
The name benanderi was proposed for C. annulatella Nylander sensu Benander (1939 : 96, pl. 5,
figs 74, 74a, pl. 6, fig. 84).
83
The identity of Phalaena (Tinea) vestianella Linnaeus, 1758
Coleophora vestianella (Linnaeus, 1758) comb. n.
(Fig. 3)
Phalaena (Tinea) vestianella Linnaeus, 1758, Syst. Nat. (edn 10) 1: 536. LECTOTYPE 63, [SwEDEN]
(genitalia slide no. 715, Sattler; Linnean Society, London), here designated [examined].
Phalaena (Tinea) vestianella Linnaeus; Linnaeus, 1761, Fauna suec. (edn 2) : 365.
Phalaena (Tinea) vestianella Linnaeus; Linnaeus, 1767, Syst. Nat. (edn 12) 1 (2) : 888.
Ornix laripennella Zetterstedt, [1839], Insecta lapponica: 1011. Syntype(s) 3, Norway: Dovrefjell
(‘alpe Dowre’) (Boheman) (? Naturhistoriska Riksmuseum, Stockholm) [not examined]. Syn. n.
Coleophora annulatella Nylander, 1848, in Tengstrém, Notis. Sdllsk. Faun. Fl. fenn. Férh. 1 : 143. Syntypes
3, 9, FINLAND: Helsinki (‘Helsingfors’), vii. (Nylander) (Zoological Museum, Helsinki) [not examined].
Syn. n.
Coleophora tengstromella Doubleday, 1859, Zoologist syn. List. Br. Butterfl. & Moths (edn 2) : 33. [Objec-
tive replacement name for C. annulatella Nylander, 1848.] Syn. n.
Coleophora annulatella Nylander; Wocke, 1861, in Staudinger & Wocke, Cat. Lepid. Eur. : 123. [With
laripennella Zetterstedt in synonymy. ]
Coleophora laripennella (Zetterstedt); Wocke, 1871, in Staudinger & Wocke, Cat. Lepid. eur. Faunen-
gebiets : 318. [With annulatella Nylander in synonymy. ]
Coleophora laripennella (Zetterstedt); Rebel, 1901, in Staudinger & Rebel, Cat. Lepid. palaearctischen
Faunengebietes 2 : 200.
Eupista laripennella (Zetterstedt); Pierce & Metcalfe, 1935, Genitalia Tineid Families Lepid. Br. Islands : 66,
pl. 40, figs (3, 9).
Coleophora laripennella (Zetterstedt); Benander, 1939, Opusc. ent. 4: 94, pl. 5, figs 72, 72a (3).
Eupista laripennella (Zetterstedt); Toll, 1953, Mater. Fizjogr. Kraju 32 : 182, pl. 15, fig. 125 (3), pl. 29,
fig. 281 (9).
Coleophora laripennella (Zetterstedt); Bradley, 1972, Handbk Ident. Br. Insects 11 (2): 16.
Coleophora laripennella (Zetterstedt); Patzak, 1974, Beitr. Ent. 24: 265, colour fig. 33 (fore wing), figs
203 (3), 337 (9).
Coleophora laripennella (Zetterstedt); Karsholt & Nielsen, 1976a, Syst. Fortegnelse over Danmarks
Sommerfugle : 28.
In 1758 Linnaeus described Phalaena Tinea vestianella as follows:
‘P. Tinea alis cinereis: apicibus adscendentibus plumosis, palpis recurvis. Habitat in Vestimentis,
quae destruit.’
He placed it after cynosbatella (Tortricidae) and before tapetzella, pellionella (Tineidae) and
sarcitrella (Oecophoridae). In 1761 he placed it after tapetzella, pellionella and sarcitrella and
before pomonella (Tortricidae), adding a more detailed description which reads as follows:
‘PH. TINEA vestianella alis cinereis costa albis, apicibus adscendentibus plumosis, palpis recurvatis.
Habitat in Vestimentis, quae aestate rodit & destruit more praecedentium.
DESCR. Alae superiores lanceolatae, margine crassiore niveo, glaucae atomis nigris. Jnferiores subu-
latae, plumbei coloris, omnes latissime ciliatae. Palpi prominentes, recurvi. Antennae breviores, albo
annulatae.’
Linnaeus’s description and placement of vestianella in 1767 differs little from that of 1761.
In the Linnaean collection there are three identical specimens (1 3g, 1 2; 1 of undetermined sex,
without abdomen), the 3 being associated with the label ‘vestianella’ in Linnaeus’s handwriting.
An additional label ‘vestianella 888’ was subsequently added by Sir James Edward Smith, who
acquired the collection after the death of Linnaeus. A detailed examination of these specimens
has shown them to be the species hitherto known as Coleophora laripennella (Zetterstedt, [1839)).
The specimens agree with the 1758 description by their grey wings and recurved palpi but dis-
agree by their biology and down-turned apices of the wings. Additional descriptive material
added by Linnaeus in 1761 and repeated in 1767 clearly applies to the specimens in his collection
or would have applied when they were fresh: lanceolate, bluish grey forewings with black specks
and white costa; awl-shaped, grey hind wings; long cilia on all wings; prominent, recurved palpi;
white-ringed antennae.
84
If, as we believe, the specimens in the Linnaean collection are authentic and are those upon
which the descriptions were based, Linnaeus’s account of the biology is incorrect. In his descrip-
tion of the biology in 1761 (slightly more detailed than that of 1758) he stated that it lives in cloth-
ing, which it nibbles in summer and destroys in the same way as the preceding species (tapetzella,
pellionella, sarcitrella). This description is sufficiently vague that it can be applied to several
species of clothes moths and even to the oecophorids Endrosis sarcitrella (L.) and Hofmannophila
pseudospretella (Stainton). It has been suggested that vestianella might be the species currently
known as Tineola bisselliella (Hummel) because the latter is so common that Linnaeus should
have known and described it. While the biology might apply to bisselliella the brief diagnosis of
the moth in 1758 and the more detailed descriptions in 1761 and 1767 clearly do not apply, for
example, the grey wings and recurved palpi (1758) and the shape, colour and markings of the
wings as well as the ringed antennae (1761; 1767).
All evidence seems to indicate that the descriptions of the moth and of the biology apply to two
different species. If vestianella is a clothes moth then the descriptions of the moth are incorrect
and the specimens in the Linnaean collection are not authentic. However, if it is the species we
know as Coleophora laripennella (Zetterstedt) then the accounts of the biology are erroneous.
We believe that the identity of this species should, in agreement with normal practice, be based
primarily on the description of the moth and on the specimens in the Linnaean collection. Such
important evidence cannot be rejected in favour of an account of the biology, the authenticity of
which is dubious. It should be noted that erroneous biological data are not uncommon in Lin-
naeus’s works, for example, Solidago as food-plant for Heodes virgaureae, Spiraea for Zygaena
filipendulae and Lonicera for Plutella xylostella.
Past authors have interpreted vestianella in different ways and its true identity has hitherto
remained unknown.
Clerck (1759 : pl. 11, fig. 11) illustrated under the name vestianella a species currently known
as Pyrausta cespitalis ([Denis & Schiffermiiller], 1775) (Pyralidae). Stephens (1834 : 344) applied
the name vestianella to the species currently known as Monopis rusticella (Hiibner, 1796) (Tineidae)
whereas Werneburg (1864 : 232, 273) considered that it might be /acteella [Denis & Schiffermiller]
[=Endrosis sarcitrella (Linnaeus, 1758)] (Oecophoridae). More recently Bradley (1972: 8)
considered that vestianella might be conspecific with Tineola bisselliella (Hummel, 1823) (Tinei-
dae).
The above designation of the lectotype places vestianella as a senior subjective synonym of
Coleophora laripennella (Zetterstedt, [1839]). Because of the confusion regarding the names
sternipennella, flavaginella, punctipennella, muehligiella, moeniacella, benanderi, laripennella and
annulatella (pp. 81-84), the introduction of the name vestianella into this group is not disruptive
but, on the contrary, introduces an element of stability; being a Linnaean name it cannot in
future be affected by any newly discovered synonymy.
The removal of the name vestianella from the Tineidae (and Oecophoridae) is also in the
interest of stability and ensures that in future it cannot threaten the well-known names of
species of economic importance.
The syntypes of annulatella were examined and the synonymy with C. l/aripennella (Zetterstedt)
was established by Kanerva (1941 : 119).
Doubleday proposed the name tengstromella ({tengstroemella auctorum, incorrect subsequent
spelling) as an objective replacement name for Coleophora annulatella Nylander, 1848, which he
erroneously considered to be preoccupied by Cerostoma annulatella Curtis, 1832, Br. Ent., 9,
no. 420.
Alphabetical catalogue of the family-group names
tCASASINI Capuse, 1971 (Sattler & Tremewan, 1974 : 198).
Incorrect original formation of the tribe name based on Casas Wallengren, 1881.
See also: Casini Capuse, 1971.
CASIGNETINA Falkovitsh, 1972; Capuse, 1973, Taxon. Famille Coleophoridae : 8; 1975, Fragm. ent.
11 : 49.
85
Type-genus: Casigneta Wallengren, 1881, nom. praeocc. [= Casignetella Strand, 1928].
Originally proposed as a tribe name Casignetini Falkovitsh, 1972, which is invalid as a family-group
name as its nominal type-genus is a junior homonym (nt. Code zool. Nom., Article 39) (see Sattler &
Tremewan, 1974 : 198). No replacement name was proposed as we considered Casignetini Falkovitsh
to be a junior subjective synonym of Coleophoridae Hiibner, [1825]. Subsequently used as a subtribe
name; again no replacement name is proposed as Casignetina Falkovitsh is currently considered to be
a junior subjective synonym of Coleophoridae Hiibner, [1825].
CASINI Capuse, 1971; Capuse, 1973, Taxon. Famille Coleophoridae : 6.
Type-genus: Casas Wallengren, 1881.
Originally proposed as ‘Casasini’, which is an incorrect formation of the tribe name based on Casas
Wallengren, 1881; subsequently emended to Casini (Capuse, 1973 : 6; 1975 : 2).
In view of the difficulties we have experienced with regard to the correct formation of the family-
group name based on Casas, it might be appropriate to comment here on Article 29 of the Code. Few
taxonomists have a thorough knowledge of the classical languages and in this respect most of us find it
difficult or impossible to use the Code without seeking advice. In our opinion any taxonomist should
be able to use the Code without the assistance of classical scholars; we therefore suggest that the
relevant rules of the Code be amended to recommendations.
COLEOPHORIDAE Hiibner, [1825] (Sattler & Tremewan, 1974 : 198).
In 1974 we based the name Coleophoridae on Coleophorae Hiibner, [1825], as, in our opinion,
Hiibner’s ‘stirps’ is the equivalent of the family as understood by current authors. In coming to this
decision we were guided by manuscript notes of Durrant in the Walsingham copy of Hiibner’s Ver-
zeichniss and by Hemming (1937 : 19). According to Durrant, Hiibner’s ‘stirps’ is the equivalent of the
family while Hemming considered it to be the subfamily. However, Hiibner’s ‘stirps’ names have not
been generally accepted as family-group names, and Franclemont (in Hemming, 1957 : 312) stated
“Whatever importance Hiibner attached to the category ‘‘Stirps’’ he certainly did not treat it as a taxon
belonging to the family-group as currently understood.’
It has also been suggested to us that the family-group name Coleophoridae should not be based on
Coleophorae Hiibner, [1825] : 426, as the type-genus Coleophora Hiibner, 1822, is not included in the
stirps Coleophorae. The Jnternational Code of Zoological Nomenclature, Article 11(e), specifies that ‘A
family-group name must, when first published, be based on the name then valid for a contained genus,
.... Our interpretation of Article 11(e) was that the name Coleophorae is clearly based on Coleophora
which is therefore automatically contained in the stirps. We believe that the Code would have used the
term ‘included genus’ (rather than ‘contained genus’) to specify that the type-genus must be included
by name.
If our interpretations are rejected the family-group name must be attributed to Bruand, 1850 : 54
(Sattler & Tremewan, 1974 : 198).
{FALKOVITSHINAE Capuse, 1972; Capuse, 1973, Taxon. Famille Coleophoridae : 5.
Incorrect subsequent formation of the subfamily name based on Falkovitshia Capuse, 1972 (see
Sattler & Tremewan, 1974: 199).
GONIODOMINA Capuse, 1973, Taxon. Famille Coleophoridae : 8; 1975, Fragm. ent. 11: 59.
Type-genus: Goniodoma Zeller, 1849.
Originally proposed as a subtribe name; currently considered to be a junior subjective synonym
(syn. n.) of Coleophoridae Hiibner, [1825].
HAPLOPTILIINA Barnes & McDunnough, 1917; Capuse, 1975, Fragm. ent. 11: 2.
Type-genus: Haploptilia Hiibner, [1825].
Originally proposed as ‘Haploptilidae’, which is an incorrect formation of the family name based
on Haploptilia Hiibner, [1825]; subsequently used as a subtribe name (in the incorrect formation
Haploptilina) (Capuse, 1973 : 12) ; finally emended to Haploptiliina (Capuse, 1975 : 2). Currently
considered to be a junior subjective synonym of Coleophoridae Hiibner, [1825] (see also Sattler &
Tremewan, 1974 : 199).
{HAPLOPTILINA Barnes & McDunnough, 1917; Capuse, 1973, Taxon. Famille Coleophoridae : 12.
Incorrect formation of the subtribe name based on Haploptilia Hiibner, [1825]. See also Sattler &
Tremewan, 1974 : 199, Haploptilidae.
Alphabetical catalogue of the genus-group names
ABARASCHIA Capuse, 1973, Taxon. Famille Coleophoridae : 20; 1975, Fragm. ent. 11: 24.
Type-species: Coleophora pagmana Toll, 1962, Acta zool. cracov. 7: 679, pl. 11K, fig. 119, pl. 7F,
fig. 58, pl. 34A, fig. 135, pl. 15M, fig. 102, by original designation and monotypy.
86
Originally proposed as a subgenus of Baraschia Capuse, 1973; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
AMSELGHIA Capuse, 1973, Taxon. Famille Coleophoridae : 18; 1975, Fragm. ent. 11 : 38.
Type-species : Coleophora fringillella Zeller, 1839, Isis, Leipzig 1839 : 208, by original designation and
monotypy.
The type-species was cited by Capuse as ‘fringilella’ which is an incorrect subsequent spelling of
fringillella Zeller, 1839.
Originally proposed as a subgenus of Amseliphora Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
AMSELIPHORA Capuse, 1971 (Sattler & Tremewan, 1974 : 200).
See also: Amselghia Capuse, 1973; Ardania Capuse, 1973.
APISTA Hiibner, [1825] (Sattler & Tremewan, 1974 : 200).
See also: Metapista Capuse, 1973.
ARDANIA Capuse, 1973, Taxon. Famille Coleophoridae : 18; 1975, Fragm. ent. 11: 38.
Type-species: Coleophora bilineatella Zeller, 1849, Linn. ent. 4: 198, 272, by original designation and
monotypy.
Originally proposed as a subgenus of Amseliphora Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
ASCLERIDUCTIA Capuse, 1973, Taxon. Famille Coleophoridae : 19; 1975, Fragm. ent. 11: 30.
Type-species: Coleophora lithargyrinella Zeller, 1849, Linn. ent. 4: 200, 394, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
AURELIANIA Capuse, 1971, Recherches morph. syst. Famille Coleophoridae : 65.
Type-species: Coleophora flavaginella Lienig & Zeller sensu Capuse, 1971 [= Coleophora benanderi
Kanerva, 1941, Suomen hyént. Aikak. 7: 120], by original designation.
In our previous catalogue (1974 : 201) we erroneously considered the type-species of Aureliania,
namely Coleophora flavaginella Lienig & Zeller sensu Capuse, 1971, to be synonymous with Coleophora
annulatella Nylander, 1848. The examination of type-material of annulatella by Kanerva (1941 : 119)
has shown it to be a synonym of Coleophora laripennella (Zetterstedt, [1839]). For annulatella sensu
auctorum Kanerva proposed the name benanderi. As noted earlier (Sattler & Tremewan, 1974 : 201)
flavaginella Lienig & Zeller is a junior subjective synonym of sternipennella Zetterstedt. The synonymy
is expressed in detail on pp. 81-82.
Under the /nt. Code zool. Nom., Article 70(a), the case of a misidentified type-species has to be
referred to the International Commission on Zoological Nomenclature. We suggest that the Commis-
sion be asked to designate as the type-species of Aureliania the nominal species actually involved,
namely Coleophora benanderi Kanerva, 1941 (Int. Code zool. Nom., Article 70(a)(i)).
The change in the name of the type-species does not affect the synonymy of the genus Aureliania,
which is still considered to be a junior subjective synonym of Coleophora Hiibner, 1822.
See also: Ductispira Capuse, 1974; Ecebalia Capuse, 1973; Nosyrislia Capuse, 1973.
BACESCUIA Capuse, 1971, Recherches morph. syst. Famille Coleophoridae : 65.
Type-species: Coleophora moeniacella Stainton sensu Capuse, 1971 [= Coleophora suaedivora Mey-
rick, [1928], Revised Handbk Br. Lepid.: 763], by original designation and monotypy.
Under the Int. Code zool. Nom., Article 70(a), the case of a misidentified type-species has to be
referred to the International Commission on Zoological Nomenclature. We suggest that the Com-
mission be asked to designate as the type-species of Bacescuia the nominal species actually involved,
namely Coleophora suaedivora Meyrick, [1928] (Int. Code zool. Nom., Article 70(a)(i)).
C. moeniacella Stainton [= muehligiella Stainton] is currently considered to be a junior subjective
synonym of C. sternipennella (Zetterstedt, [1839]) (see p. 81).
BARASCHIA Capuse, 1973, Taxon. Famille Coleophoridae : 20; 1975, Fragm. ent. 11: 24.
Type-species: Coleophora paradoxella Toll, 1961, Annis zool. Warsz. 19 : 215, text-figs 14-18, by
original designation.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
See also: Abaraschia Capuse, 1973.
BENANDERPIA Capuse, 1973, Taxon. Famille Coleophoridae : 10; 1975, Fragm. ent. 11: 54. <
Type-species: Coleophora adspersella Benander, 1939, Opusc. ent. 4: 93, pl. 5, fig. 70, by original
designation and monotypy.
87
The type-species was cited by Capuse as ‘adspereslla’, which is an incorrect subsequent spelling of
C. adspersella Benander, 1939.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
{CALARITANIA Mariani, 1943, G. Sci. nat. econ. Palermo 42 (3): 188 (nomen nudum). (Sattler &
Tremewan, 1974 : 202, t Calaritania Grandi.)
{ Calaritania Mariani was not accompanied by a description, indication or available nominal species;
the only included nominal species, tsardiniella, is a nomen nudum. Mariani attributed the names
Calaritania and sardiniella to Amsel, who, however, has never made them nomenclaturally available
(Amsel, in litt.).
CALCOMARGINIA Capuse, 1973, Taxon. Famille Coleophoridae : 17; 1975, Fragm. ent. 11: 38.
Type-species: Ornix ballotella Fischer von Roeslerstamm, 1839, Abbildungen Bericht. Ergdanzung
Schmettkde : 154, pl. 58, figs a-k, by original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
{CALEOPHORA Capuse, 1973, Taxon. Famille Coleophoridae : 11 [under Hamuliella]. Incorrect subse-
quent spelling of Coleophora Hiibner, 1822.
COLEOPHORA Hiibner, 1822 (Sattler & Tremewan, 1974 : 203).
See also: Abaraschia Capuse, 1973; Amselghia Capuse, 1973; Ardania Capuse, 1973, Ascleriductia
Capuse, 1973; Baraschia Capuse, 1973; Benanderpia Capuse, 1973; Calcomarginia Capuse, 1973;
¢Caleophora Capuse, 1973; tCorothropoea Capuse, 1973; Cornulivalvulia Capuse, 1973; Ductispira
Capuse, 1974; Dumitrescumia Capuse, 1973; Ecebalia Capuse, 1973; Globulia Capuse, 1973; Hamuliella
Capuse, 1973; Helvalbia Capuse, 1973; Kasyfia Capuse, 1973; Klimeschjosefia Capuse, 1975; Kuznet-
zovvlia Capuse, 1973; Latisacculia Capuse, 1973; Longibacillia Capuse, 1973; Lucidaesia Capuse, 1973;
Luzulina Falkovitsh, 1972; Lvaria Capuse, 1973; Membrania Capuse, 1973; Metapista Capuse, 1973;
Multicoloria Capuse, 1973; Neugenvia Capuse, 1973; Nosyrislia Capuse, 1973; t Ortohgraphis Capuse,
1973; Oudejansia Capuse, 1973; Paravalvulia Capuse, 1973; Patzakia Capuse, 1973; Perygridia Falko-
vitsh, 1972; Postvinculia Capuse, 1973; Proglaseria Capuse, 1973; Quadratia Capuse, 1973; Rhamnia
Capuse, 1973; Sacculia Capuse, 1973; Scleriductia Capuse, 1973; Tollsia Capuse, 1973; Tuberculia
Capuse, 1973; Ulna Capuse, 1973.
CORETHROPOEA Falkovitsh, 1972 (Sattler & Tremewan, 1974 : 203)..
See also: { Corothropoea Capuse, 1973.
{COROTHROPOEA CaApuse, 1973, Taxon. Famille Coleophoridae : 20.
Incorrect subsequent spelling of Corethropoea Falkovitsh, 1972.
CORNULIVALVULIA Capuse, 1973, Taxon. Famille Coleophoridae : 9; 1975, Fragm. ent. 11: 49.
Type-species: Coleophora vicinella Zeller, 1849, Linn. ent. 4: 198, 251, by original designation and
monotypy.
Junior subjective synonym of Multicoloria Capuse, 1973 (Falkovitsh, 1974a : 194); currently con-
sidered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
CORYTHANGELA Meyrick, 1897 (Sattler & Tremewan, 1974 : 203).
See also: { Corythangella Capuse, 1973.
{CORYTHANGELLA CaAapuse, 1973, Taxon. Famille Coleophoridae : 22.
Incorrect subsequent spelling of Corythangela Meyrick, 1897.
DUCTISPIRA Capuse, 1974, Beitr. naturk. Forsch. Siidw Dtl. 33 : 188.
Type-species: Coleophora unistriella Caradja, 1920, Dt. ent. Z. Iris 34: 152, by original designation
and monotypy.
Originally proposed as a subgenus of Aureliania Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
DUMITRESCUMIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 8.
Type-species: Coleophora cecidophorella Oudejans, 1972, Ent. Ber., Amst. 32: 120, by original
designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822. :
ECEBALIA Capuse, 1973, Taxon. Famille Coleophoridae : 11; 1975, Fragm. ent. 11 : 53.
Type-species: Ornix laripennella Zetterstedt, [1839], Insecta lapponica : 1011, by original designation
and monotypy.
For synonymy of C. lJaripennella (Zetterstedt, [1839]) see p. 84.
Originally proposed as a subgenus of Aureliania Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
88
GLASERIA Capuse, 1971 (Sattler & Tremewan, 1974 : 204).
See also: Proglaseria Capuse, 1973. .
GLOBULIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 11.
Type-species: Coleophora cornuta Heinemann & Wocke, [1876], in Heinemann, Schmett. Dtl.
Schweiz (2) 2 (2) : 539, by original designation and monotypy.
Capuse cited the type-species as ‘Coleophora cornuta Stt., 1859’, which is a nomen nudum. The species
was first made nomenclaturally available by Heinemann & Wocke, [1876]; the synonymy is expressed
as follows:
Coleophora cornuta Heinemann & Wocke, [1876], in Heinemann, Schmett. Dtl. Schweiz (2)
2. (2) 2539.
Coleophora cornuta Stainton, 1859, Nat. Hist. Tineina 4: 14, 15, 32, 33. Nomen nudum.
Coleophora cornuta Wocke, 1861, in Staudinger & Wocke, Cat. Lepid. Eur. : 121. Nomen nudum.
Coleophora cornuta Wocke, 1871, in Staudinger & Wocke, Cat. Lepid. eur. Faunengebiets : 313.
Nomen nudum.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
HAMULIELLA Capuse, 1973, Taxon. Famille Coleophoridae : 11; 1975, Fragm. ent. 11: 53.
Type-species: Coleophora otitae Zeller, 1839, Isis, Leipzig 1839 : 207, by original designation and
monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
HELVALBIA Capuse, 1973, Taxon. Famille Coleophoridae : 19; 1975, Fragm. ent. 11 : 32.
Type-species: Porrectaria lineolea Haworth, 1828, Lepid. Br. : 534, by original designation and
monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
KASYFIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 16.
Type-species: Ornix binderella Kollar, 1832, Beitr. Landesk. Ost. Enns 2 : 99, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
KLIMESCHJA Capuse, 1971 (Sattler & Tremewan, 1974 : 205).
See also: Klimeschjosefia Capuse, 1975.
KLIMESCHJOSEFIA Capuse, 1975, Fragm. ent. 11 : 35 (objective replacement name for Klimeschja
Capuse, 1971).
Type-species: Coleophora oriolella Zeller, 1849, Linn. ent. 4: 198, 258, by original designation for
and monotypy of Klimeschja Capuse, 1971.
Unnecessary objective replacement name for Klimeschja Capuse, 1971, which is not a junior homonym
of Klimeschia Amsel, 1938 (Lepidoptera : Douglasiidae). Capuse was aware of the Code, Article
56(a) (one-letter difference) and stated: ‘Nous pensons que les lettres i—j entrent aussi dans cette régle.’
In spite of this he continued: ‘Contrairement, nous proposons pour notre genre le nom Klimeschjosefia,
.... The latter statement is not clear to us. Did Capuse propose a replacement name in spite of Klime-
schja being valid under the Code or had he doubts about the one-letter difference i-j and therefore
proposed a conditional replacement name? If the former was intended then it is an unnecessary replace-
ment name; if the latter was intended, then it is a conditionally proposed name and unavailable under
the Code, Article 15.
KUZNETZOVVLIA Capuse, 1973, Taxon. Famille Coleophoridae : 19; 1975, Fragm. ent. 11: 25.
Type-species: Coleophora solidaginella Staudinger, 1859, Stettin. ent. Ztg 20: 254, by original
designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
_ LATISACCULIA Capuse, 1973, Taxon. Famille Coleophoridae : 18; 1975, Fragm. ent. 11: 32.
Type-species: Coleophora crocinella Tengstrém, 1848, Notis. Sdllsk. Faun. Fl. fenn. Férh. 1: 140, by
original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
LONGIBACILLIA Capuse, 1973, Taxon. Famille Coleophoridae : 20; 1975, Fragm. ent. 11: 23.
Type-species: Coleophora fergana Toll, 1961, Annls zool. Warsz. 19 : 211, text-figs 5-9, by original
designation and monotypy.
89
Originally proposed as a subgenus of Membrania Capuse, 1973; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
LUCIDAESIA Capuse, 1973, Taxon. Famille Coleophoridae : 15; 1975, Fragm. ent. 11 : 47.
Type-species: Phalaena (Tinea) frischella Linnaeus sensu Capuse, 1973 [= Ornix alcyonipennella
Kollar, 1832, Beitr. Landesk. Ost. Enns 2: 99], by original designation and monotypy.
Under the Int. Code zool. Nom., Article 70(a), the case of a misidentified type-species has to be
referred to the International Commission on Zoological Nomenclature. We suggest that the Com-
mission be asked to designate as the type-species of Lucidaesia the nominal species actually involved,
namely Coleophora alcyonipennella (Kollar, 1832) (Int. Code zool. Nom., Article 70(a)(i)).
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
LUZULINA Falkovitsh, 1972 (Sattler & Tremewan, 1974 : 205).
Originally proposed as a subgenus of Perygra Falkovitsh, 1972; subsequently considered as a genus
(Capuse, 1973 : 11); currently considered to be a junior subjective synonym of Coleophora Hiibner,
1822 (Sattler & Tremewan, 1974 : 205).
LVARIA Capuse, 1973, Taxon. Famille Coleophoridae : 9; 1975, Fragm. ent. 11: 57.
Type-species: Coleophora lassella Staudinger, 1859, Stettin. ent. Ztg 20 : 255, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
MEMBRANIA Capuse, 1973, Taxon. Famille Coleophoridae : 19; 1975, Fragm. ent. 11: 22, 23.
Type-species: Coleophora calycotomella Stainton, 1869, Tineina sth. Eur. : 225, by original desig-
nation.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
See also: Longibacillia Capuse, 1973.
METAPISTA Capuse, 1973, Taxon. Famille Coleophoridae : 17; 1975, Fragm. ent. 11: 41.
Type-species: Coleophora stramentella Zeller, 1849, Linn. ent. 4: 198, 274, by original designation
and monotypy.
Originally proposed as a subgenus of Apista Hiibner, [1825]; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
MULTICOLORIA Capuse, 1973, Taxon. Famille Coleophoridae : 17; 1975, Fragm. ent. 11: 43.
Type-species: Coleophora ditella Zeller, 1849, Linn. ent. 4: 198, 247, by original designation and
monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
See also: Cornulivalvulia Capuse, 1973.
NEUGENVIA Capuse, 1973, Taxon. Famille Coleophoridae : 10; 1975, Fragm. ent. 11 : 54.
Type-species: Eupista vlachi Toll, 1953, Mater. Fizjogr. Kraju 32: 200, pl. 17, fig. 141, pl. 30,
fig. 293, pl. 34, fig. 46, by original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
NOSYRISLIA Capuse, 1973, Taxon. Famille Coleophoridae : 11; 1975, Fragm. ent. 11: 52.
Type-species: Coleophora linosyris Hering, 1937, Mitt. zool. Mus. Berl. 22 : 280, text-figs, by original
designation and monotypy.
Originally proposed as a subgenus of Aureliania Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
ORTHOGRAPHIS Falkovitsh, 1972 (Sattler & Tremewan, 1974 : 206).
See also: { Ortohgraphis Capuse, 1973.
tORTOHGRAPHIS Capuse, 1973, Taxon. Famille Coleophoridae : 17.
Incorrect subsequent spelling of Orthographis Falkovitsh, 1972.
OUDEJANSIA Capuse, 1973, Taxon. Famille Coleophoridae : 20; 1975, Fragm. ent. 11: 21.
Type-species: Coleophora obviella Rebel, 1914, Verh. zool.-bot. Ges. Wien 64: (178), by original
designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
PARAVALVULIA Capuse, 1973, Taxon. Famille Coleophoridae : 13; 1975, Fragm. ent. 11: 5.
Type-species: Coleophora spiraeella Rebel, 1916, Verh. zool.-bot. Ges. Wien 66: (15), by original
designation and monotypy.
90
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
PATZAKIA Capuse, 1973, Taxon. Famille Coleophoridae : 10; 1975, Fragm. ent. 11: 56.
Type-species: Coleophora silenella Herrich-Schiaffer, 1855, Syst. Bearb. Schmett. Eur. 5 : 227 [key],
252; 1854, ibidem 5, pl. 113, fig. 920d [as dianthi, non-binominal; misidentification], by original desig-
nation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
PERYGRA Falkovitsh, 1972 (Sattler & Tremewan, 1974 : 206).
See also: Luzulina Falkovitsh, 1972; Perygridia Falkovitsh, 1972.
PERYGRIDIA Falkovitsh, 1972 (Sattler & Tremewan, 1974 : 206).
Originally proposed as a subgenus of Perygra Falkovitsh, 1972; subsequently considered as a genus
(Capuse, 1973 : 10); currently considered to be a junior subjective synonym of Coleophora Hibner,
1822 (Sattler & Tremewan, 1974 : 206).
POSTVINCULIA Capuse, 1973, Taxon. Famille Coleophoridae : 13; 1975, Fragm. ent. 11: 4.
Type-species: Coleophora lutipennella Zeller, 1838, Isis, Leipzig 1838 : 713, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
PROGLASERIA Capuse, 1973, Taxon. Famille Coleophoridae : 18; 1975, Fragm. ent. 11 : 27.
Type-species: Coleophora laticostella Mann, 1859, Wien. ent. Monatschr. 3 : 175, by original desig-
nation and monotypy.
Originally proposed as a subgenus of Glaseria Capuse, 1971; currently considered to be a junior
subjective synonym (syn. n., see pp. 74-75) of Coleophora Hiibner, 1822.
QUADRATIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 13.
Type-species: Coleophora fuscocuprella Herrich-Schaffer, 1854, Syst. Bearb. Schmett. Eur. 5: 220
[key]; 1854, ibidem5, pl. 113, fig. 920f [as ‘Fuscocupr.’, non-binominal]; 1855, ibidem 5 : 230, by original
designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
RHAMNIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 18.
Type-species: Coleophora ahenella Heinemann, [1876], Schmett. Dtl. Schweiz (2) 2 (2) : 546, by
original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
SACCULIA Capuse, 1973, Taxon. Famille Coleophoridae : 13; 1975, Fragm. ent. 11: 7.
Type-species: Coleophora excellens Toll, 1952, Bull. Soc. ent. Mulhouse 1952: 20, pl. 1, fig. 6, by
original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
SCLERIDUCTIA Capuse, 1973, Taxon. Famille Coleophoridae : 20; 1975, Fragm. ent. 11: 28.
Type-species: Coleophora ochripennella Zeller, 1849, Linn. ent. 4: 200, 408, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
TOLLSIA Capuse, 1973, Taxon. Famille Coleophoridae : 21; 1975, Fragm. ent. 11: 15.
Type-species: Coleophora hornigi Toll, 1952, Bull. Soc. ent. Mulhouse 1952: 18, pl. 1, fig. 3, by
original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
C. hornigi Toll, 1952, is currently considered to be a junior subjective synonym of Phalaena (Tinea)
violacea Strém, 1783, Nye Saml. K. dansk. Vid. Selsk. Skr. 2: 88 (Karsholt & Nielsen, 19765 : 245).
TUBERCULIA Capuse, 1973, Taxon. Famille Coleophoridae : 17; 1975, Fragm. ent. 11: 9.
Type-species: Coleophora albitarsella Zeller, 1849, Linn. ent. 4: 200, 378, by original designation
and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
ULNA Capuse, 1973, Taxon. Famille Coleophoridae : 9; 1975, Fragm. ent. 11: 58.
91
Type-species: Coleophora saponariella Heeger, 1848, Isis, Leipzig 1848 : 342, pl. 6, figs A-N, by
original designation and monotypy.
Currently considered to be a junior subjective synonym (syn. n., see pp. 74-75) of Coleophora
Hiibner, 1822.
References
Benander, P. 1938-39. Die Coleophoriden Schwedens. Opusc. ent. 3 : 107-124 (1938); ibidem 4 : 30-110,
pls 1-8 (1939).
—— 1940. Revision von Zetterstedts lapplandischen Microlepidoptera. Opusc. ent. 5 : 49-65, pl. 1.
Bradley, J. D. 1962. The identity of certain species of Coleophoridae (Lepidoptera). Entomologist’s Gaz.
13 : 178-184, figs 1-3.
—— 1972. In Kloet, G. S. & Hincks, W. D., A check list of British insects. Second edition (revised).
Part 2: Lepidoptera. Handbk Ident. Br. Insects 11 (2) : i-viii, 1-153.
Bruand, Th. 1850. Catalogue systématique et synonymique des lépidoptéres du département du Doubs.
Mém. Soc. Emul. Doubs (1) 3 (3) : 23-68.
Capuse, I. 1971. Recherches morphologiques et systématiques sur la famille des Coleophoridae (Lepidoptera).
66 pp., 46 pls. Bucarest.
—— 1973. Sur la taxonomie de la famille des Coleophoridae (Clés de détermination des taxa superspéci-
fiques). 24 pp. Bucaresti.
—— 1974. Beitrage zum Studium der Familie Coleophoridae, IX (Lepidoptera). Uber die von A. Caradja
beschriebenen Taxa der Gattung Coleophora. Beitr. naturk. Forsch. SiidwDtl. 33 : 179-195, pl. 1,
figs A-F, text-figs 1-11.
—— 1975. Complément systématique a la famille des Coleophoridae (Lepidoptera). Fragm. ent. 11 : 1-64,
text-figs 1-27.
Clerck, C. 1759-64. Icones insectorum rariorum. [viii], [iii], [iii] pp., 55 pls. Holmiae.
Falkovitsh, M. I. 1972. New genera of palaearctic casebearers (Lepidoptera, Coleophoridae). [In Russian.]
Ent. Obozr. 51 : 369-386, text-figs 1-32.
—— 1973. Contribution to the knowledge of casebearers (Lepidoptera, Coleophoridae) of the Kisilkum
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1974a. A new species of the genus Multicoloria Cap. (Lepidoptera, Coleophoridae) living on
Eremosparton. [In Russian.] Ent. Obozr. 53 : 194-195, text-fig.
1974b. A new species of the genus Multicoloria Cap. (Lepidoptera, Coleophoridae) living on
Eremosparton. [English translation of Falkovitsh, 1974a.] Ent. Rev., Wash. 53(1) : 138-139, text-fig.
Hemming, F. 1937. Hiibner. 1. xxxiv, 605 pp., frontispiece. London.
—— 1957. Family-group-name problems involved in the decision under the vote taken on Voting
Paper V.P. (54) 43 to validate as of subgeneric status as from Linnaeus, 1758, certain names in the
Order Lepidoptera (Class Insecta) originally published as terms for groups of species within the genus
“‘Phalaena” Linnaeus, 1758 or 1767. Opin. Decl. Int. Commn zool. Nom. 15 : 302-318.
Kanerva, N. 1941. Mitteilungen iiber die Coleophoriden (Lep.) Finnlands. I-II. Suomen hyént. Aikak.
7: 117-127, text-figs 1-5.
Karsholt, O. & Nielsen, E. S. 1976a. Systematisk fortegnelse over Danmarks sommerfugle. 128 pp. Klam-
penborg.
—— 1976b. Notes on some Lepidoptera described by Linnaeus, Fabricius, and Strém. Ent. scand.
7: 241-251, text-fig. 1.
Meyrick, E. [1928]. A revised handbook of British Lepidoptera. 914 pp., text-figs. London.
Patzak, H. 1974. Beitrage zur Insektenfauna der DDR: Lepidoptera—Coleophoridae. Beitr. Ent. 24 : 153-
278, colour figs 1-34, text-figs 1-363.
Pierce, F. N. & Metcalfe, J. W. 1935. The genitalia of the Tineid families of the Lepidoptera of the British
Islands. xxii, 116 pp., 68 pls. Oundle, Northants.
—— 1938. The genitalia of the British Pyrales with the Deltoids and Plumes. xiii, 69 pp., frontispiece, 29 pls.
Oundle, Northants. ,
Reznik, S. Ya. 1976. New species of the genus Multicoloria Cap. (Lepidoptera, Coleophoridae) from
USSR and adjacent countries. [In Russian.] Ent. Obozr. 55 : 648-656, text-figs 1-22.
Sattler, K. & Tremewan, W. G. 1974. A catalogue of the family-group and genus-group names of the
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Index
Synonyms and unavailable names are in /ralics. Page numbers of principal references are in bold.
Abaraschia Capuse, 76, 86
acrisella Milliére, 77
adspersella Benander, 80, 87
agenjoi Toll, 80
agnatella Toll, 78
agrianella Rebel, 78
ahenella Heinemann, 76, 91
albarracinica Toll, 79
albicosta Haworth, 77
albicostella Duponchel, 77
albidorsella Toll, 77
albitarsella Zeller, 76, 91
albostraminata Toll, 78
alcyonipennella Kollar, 90
algeriensis Toll, 77
amasiella Staudinger, 79
amethystinella Ragonot, 79
Amselghia Capuse, 78, 87
Amseliphora Capuse, 77, 87
anatolica Toll, 78
andalusiae Toll, 78
angustilinea Toll, 76
annulatella Nylander, 83, 84, 85, 87
Apista Hiibner, 78, 87
Ardania Capuse, 78, 87
arenariella Zeller, 77
arenicola Toll, 78
argentifimbriata Walsingham, 79
argyrella Herrich-Schaffer, 79
artemisicolella Bruand, 80
Ascleriductia Capuse, 77, 87
asterifoliella Klimesch, 80
asteris Miihlig, 80
astragalella Zeller, 78
atlanticolella Zerny, 79
atriplicis Meyrick, 83
atticae Capuse, 75, 79
Aureliania Capuse, 79, 81, 87
auroguttella Zeller, 81
auronitella Toll, 79
axana Hering, 80
Bacescuia Capuse, 81, 87
ballotella Fischer von Roeslerstamm, 78, 88
Baraschia Capuse, 76, 87
basimaculella Mann, 79
benanderi Kanerva, 81, 82, 83, 85, 87
Benanderpia Capuse, 80, 87
berlandella Toll, 79
bilineatella Zeller, 78, 87
bilineella Herrich-Schaffer, 77
binderella Kollar, 76, 89
binotapennella Duponchel, 81
biseriatella Staudinger, 76
biskraensis Toll, 79
bisselliella Hummel (Tineola), 85
botauripennella Toll, 80
Bourgogneja Capuse, 81
bruneosignata Toll, 77
caelebipennella Zeller, 78
Calaritania Mariani, 88
Calcomarginia Capuse, 78, 88
Caleophora Capuse, 88
calycotomella Stainton, 76, 90
canariipennella Toll, 78
Carpochena Falkovitsh, 81
Carpochenini, 74, 81
cartilaginella Christoph, 79
Casas Wallengren, 75, 85, 86
Casasini, 85, 86
Casigneta Wallengren, 86
Casignetella Strand, 86
Casignetina, 79, 85
Casignetini, 74, 75, 79, 86
Casini, 75, 85, 86
cecidophorella Oudejans, 76, 88
cespitalis Denis & Schiffermiiller (Pyrausta), 85
Characia Falkovitsh, 76
ciconiella Herrich-Schaffer, 81
cinerea Toll, 80
clypeiferella Hofmann, 81
coarctella Staudinger, 77
Coleophora Hiibner, 75, 86, 88
Coleophoridae, 74, 75, 86
Coleophorina, 76
Coleophorinae, 74, 75
Coleophorini, 74, 75
colutella Fabricius, 78
conspicuella Zeller, 78
93
conyzae Zeller, 76
coracipennella Hiibner, 75
Corethropoea Falkovitsh, 88
Cornulivalvulia Capuse, 75, 79, 88
cornuta Heinemann & Wocke, 76, 89
cornuta Stainton, 89
Corothropoea Capuse, 88
Corythangela Meyrick, 88
Corythangella Capuse, 88
cribrella Toll, 76
crocinella Tengstr6m, 77, 89
cuencella Toll, 76
cuprariella Zeller, 79
cuprifulgella Toll, 79
cynosbatella Linnaeus (Epiblema), 84
cyrniella Rebel, 79
Damophila Curtis, 79
dannehli Toll, 79
defessella Herrich-Schaffer, 77
dentiferella Toll, 80
depauperella Toll, 78
dianthivora Walsingham, 80
dichroella Toll, 76
didyma Toll, 78
discordella Zeller, 77
ditella Zeller, 78, 90
Douglasiidae, 89
dubiella Baker, 79
dubiosa Toll, 78
Ductispira Capuse, 88
Dumitrescumia Capuse, 76, 88
Ecebalia Capuse, 80, 88
echinella Staudinger, 79
erigerella Ford, 80
escalerai Toll, 78
eucera Toll, 78
excellens Toll, 75, 91
Falkovitshia Capuse, 86
Falkovitshinae, 86
femorella Walsingham, 77
fergana Toll, 76, 89
filipendulae Linnaeus (Zygaena), 85
fiorii Toll, 80
fischeri Toll, 80
flabelligerella Rebel, 79
flavaginella Lienig & Zeller, 79, 81, 82, 83, 85, 87
flavagipennella Lienig & Zeller, 81
flavipennella Herrich-Schaffer, 75
fraudulentella Toll, 77
Frederickoenigia Capuse, 75
fretella Zeller, 77
fringillella Zeller, 78, 87
frischella Linnaeus, 79
frischella Linnaeus sensu Capuse, 90
fulvosquamella Herrich-Schaffer, 80
fuscatella Toll, 77
fuscicornis Zeller, 79
fuscoaenea Toll, 79
fuscociliella Zeller, 79
fuscocuprella Herrich-Schiaffer, 76, 91
fuscostraminella Toll, 79
galatellae Hering, 80
gallipennella Hiibner, 78
gardesanella Toll, 80
gaviaepennella Toll, 80
gazella Toll, 79
genistae Stainton, 77
gilveolella Toll, 77
giraudi Toll, 78
glaseri Toll, 77
Glaseria Capuse, 76, 89
Globulia Capuse, 76, 89
gnaphalii Zeller, 80
Goniodoma Zeller, 81, 86
Goniodomina, 81, 86
gozmanyi Toll, 79
graminicolella Heinemann, 81
granulatella Zeller sensu Klimesch, 80
granulosella Staudinger, 80
griseomixta Toll, 80
gryphipennella Bouché, 75
halophylella Zimmermann, 79
haloxyli Falkovitsh, 76
Hamuliella Capuse, 80, 89
Haploptilia Hiibner, 75, 86
Haploptiliina, 75, 86
Haploptilina, 86
hartigi Toll, 77
helichrysella Krone, 79
Helvalbia Capuse, 77, 89
heringi Toll, 80
herzegoviniensis Capuse, 75, 79
hieronella Zeller, 79
honestella Toll, 79
hornigi Toll, 76, 91
hsiaolingella Toll, 80
hydrolapathella Hering, 76
imbecilla Toll, 77
insulicola Toll, 80
involucrella Chrétien, 78
Jonescumia Capuse, 81
iranella Toll, 78
jerusalemella Toll, 76
Kasyfia Capuse, 76, 89
kasyi Toll, 78
klemensiewiczi Toll, 80
Klimeschia Amsel, 89
Klimeschja Capuse, 77, 89
Klimeschjosefia Capuse, 77, 89
Klinzigedia Capuse, 81
Kuznetzovvlia Capuse, 76, 89
94
lacteella Denis & Schiffermiiller (Endrosis), 85
laripennella Zetterstedt, 80, 84, 85, 87, 88 ©
lassella Staudinger, 81, 90
laticostella Mann, 76, 91
Latisacculia Capuse, 77, 89
leucapennella Hiibner, 75
leucogrammella Herrich-Schaffer, 80
leucostoma Gerasimov, 79
leucostrigella Toll, 78
lewandowskii Toll, 80
lineariella Zeller, 80
lineata Toll, 77
lineolea Haworth, 77, 89
linosyris Hering, 80, 90
lithargyrinella Zeller, 77, 87
Longibacillia Capuse, 76, 89
Lucidaesia Capuse, 79, 90
luteolella Staudinger, 80
lutipennella Zeller, 75, 91
Luzulina Falkovitsh, 90
Lvaria Capuse, 81, 90
maeniacella auctorum, 83
mandschuriae Toll, 78
marianii Toll, 77
medelichensis Krone, 77
medicaginella Mann, 77
medicaginis Herrich-Schaffer, 78
medicagivora Toll, 79
mellechella Toll, 76
Membrania Capuse, 76, 90
mendosella Toll, 80
menephilella Toll, 79
Metapista Capuse, 78, 90
microeucera Toll, 79
microxantha Walsingham, 76
minaxella Toll, 79
moeniacella Stainton, 81, 83, 85, 87
moestella Toll, 76
mongetella Chrétien, 77
motacillella Zeller, 80
muehligella Wocke, 83
muehligiella Stainton, 81, 83, 85, 87
Multicoloria Capuse, 75, 78, 90
murciana Toll, 77
namangana Toll, 76
Neugenvia Capuse, 80, 90
niveicostella Zeller, 77
niveopictella Toll, 76
Nosyrislia Capuse, 80, 90
obliterata Toll, 80
obtectella Zeller, 76
obviella Rebel, 76, 90
ochripennella Zeller, 77, 91
ochristrigella Ragonot, 78
Oecophoridae, 84, 85
onobrychiella Zeller, 77
ononidella Milliére, 78
onosmella Brahm, 81
opacella Toll, 80
orbitella Zeller, 76
Orghidania Capuse, 75
oriolella Zeller, 77, 89
Orthographis Falkovitsh, 90
Ortohgraphis Capuse, 90
Otitae Zeller, 80, 89
Oudejansia Capuse, 76, 90
pabulella Zeller, 77
pagmana Toll, 76, 86
palumbipennella Toll, 80
paradoxella Toll, 76, 87
paragenistae Kasy, 77
paragiraudi Toll, 78
paraobviella Toll, 76
paraononidella Amsel, 78
paraspumosella Toll, 78
Paravalvulia Capuse, 75, 90
parcella Toll, 76
parenthella Toll, 80
partitella Zeller, 78
Patzakia Capuse, 80, 91
pellionella Linnaeus (Tinea), 84, 85
peribenanderi Toll, 80
perplexella Toll, 77
perserenella Rebel, 78
Perygra Falkovitsh, 91
Perygridia Falkovitsh, 91
phlomidella Christoph, 81
pirizanella Toll, 78
polonicella Zeller, 78
pomonella Linnaeus (Cydia), 84
Postvinculia Capuse, 75, 91
praecipua Walsingham, 79
praeposita Toll, 79
pratella Zeller, 80
Proglaseria Capuse, 76, 91
protecta Walsingham, 77
pseudoobviella Toll, 76
pseudoserenella Toll, 77
pseudospretella Stainton (Hofmannophila), 85
ptarmica Walsingham, 76
punctipennella Nylander, 81, 83, 85
Pyralidae, 85
Quadratia Capuse, 76, 91
qulikushella Toll, 78
radiosella Toll, 79
ramosella Zeller, 80
rebeli Gerasimov, 78
rectilineella Fischer von Roeslerstamm, 77
rectivittella Capuse, 75, 79
Rhamnia Capuse, 76, 91
riffelensis Rebel, 80
robustella Fuchs, 80
95
rudella Toll, 77 tadzhikiella Danilevski, 75
rugulosa Toll, 77 tapetzella Linnaeus (Trichophaga), 84, 85
rusticella Hiibner (Monopis), 85 tengstroemella auctorum, 85
tengstromella Doubleday, 84, 85
sabulella Toll, 80 therinella Tengstr6m, 79
Sacculia Capuse, 75, 91 Tineidae, 84, 85
santolinella Constant, 76 tolli Klimesch, 80
saponariella Heeger, 81, 92 Tollsia Capuse, 76, 91
sarcitrella Linnaeus (Endrosis), 84, 85 Tortricidae, 84
satellitella Toll, 78 tozeurensis Toll, 79
saturatella Stainton, 77 trifariella Zeller, 77
scabrida Toll, 80 trifisella Rebel, 76
schahkuhensis Toll, 76 trifolii Curtis, 79
Scleriductia Capuse, 77, 91 trigeminella Fuchs, 76
semicinerea Staudinger, 77 tristraminata Toll, 78
semistriatella Toll, 76 tritici Lindemann, 81
semistrigata Toll, 78 troglodytella Duponchel, 80
separatella Benander, 80 Tuberculia Capuse, 76, 91
silenella Herrich-Schaffer, 80, 91 turbatella Toll, 77
siliquella Constant, 77
similis Staudinger, 78 Ulna Capuse, 81, 91
simillima Staudinger, 78 unistriella Caradja, 88
solidaginella Staudinger, 76, 89 uralensis Toll, 76
spiraeella Rebel, 75, 90
spissicornis Haworth, 79
spumosella Toll, 78
squalorella Zeller, 81
squamella Constant, 77
squamosella Stainton, 80
stachi Toll, 78
sternipennella Zetterstedt, 79, 81, 82, 83, 85, 87
Stollia Capuse, 81
stramentella Zeller, 78, 90
striolatella Toll, 76
suaedivora Meyrick, 83, 87
valesianella Zeller, 79
versurella Zeller, 79
vestianella Linnaeus, 81, 82, 84, 85
vibicella Hiibner, 78
vibicigerella Zeller, 78
vicinella Zeller, 79, 88
violacea Strom, 91
virgaureae Linnaeus (Heodes), 85
virgaureae Stainton, 79
vlachi Toll, 80, 90
vulnerariae Zeller, 77
subahenella Capuse, 76 oer a
subechinella Toll, 75, 79 USSU EEL Reta
subexcellens Toll, 75 xanthoargentea Toll, 78
suboriolella Toll, 77 xanthoptera Toll, 77
subsimilis Capuse, 75, 79 xylostella Linnaeus (Plutella), 85
succursella Herrich-Schaffer, 80
sumptuosa Toll, 79 Zagulajevia Capuse, 75
supinella Ortner, 77 zonatella Toll, 78
96
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The anomalous ant-attended mealybugs (Homoptera:
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A catalogue of the type-specimens of the Scarabaeinae
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Bulletin of the | Aine :
British Museum (Naturail "Hitory)
A catalogue of the type-specimens of the
Scarabaeinae (Scarabaeidae) and the smaller
Lamellicorn families (Coleoptera) described
by G. J. Arrow
M. E. Bacchus
Entomology series Vol 37 No3 25 May 1978
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology and Zoology, and an Historical series.
Parts are published at irregular intervals as they become ready. Volumes will contain about
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1978
ISSN 0524-6431 Entomology series
Vol 37 No 3 pp 97-115
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 May 1978
A catalogue of the type-specimens of the Scarabaeinae
(Scarabaeidae) and the smaller lamellicorn families
(Coleoptera) described by G. J. Arrow ~—,_
M. E. Bacchus __ /s GENERAL “aN
Department of Entomology, British Museum (Natural History),
Cromwell Road, London SW7 5BD
\p
Contents
BYOODSIS’. 7. xs : - Sp gs ee Reg eee on ete ee BOT:
FREPOUNCHON 5 bie ee Oe ee oe at el Pee a , Mibre OF
Abbreviations . ese et : ‘Sees : ee ek FO he . 98
Acknowledgements : ee ; : bi et a OS
Catalogue of the type-specimens described by G. J; “Arrow , : ; , , i. pr BD
Scarabaeinae ; - ; F : ; : : ( ’ ‘ . . 99
Lucanidae . , : : ; z : P : : A ; ; : : . 108
Passalidae . eae : ee. : ee. 4 2 ; , : ; : 2 20
Trogidae . ; 2 , : : fan hi ; ; Shea : ; : oe A
Hybosoridae aR A ee tee, eee ee en ee 8 PP
Geotrupidae : ; : : : : : : : ; Ein ‘ wer
References... 3 he 5 , a ne “Saat: : P : a aed
Synopsis
A list is given of all the traceable type-specimens of the species and varieties of Scarabaeidae
(Scarabaeinae), Lucanidae, Passalidae, Trogidae, Hybosoridae and Geotrupidae described by G. J.
Arrow. The current valid name or generic assignment is given where appropriate. Lectotypes are
designated where necessary and where this did not require dissection of the specimens to determine the
sex.
Introduction
Gilbert Arrow was well aware of the importance of types and always labelled one specimen
(occasionally two) from a series as ‘type’. However, the vague or ambiguous nature of his
published type designations, when any were made at all, renders most of these ‘types’, except
the uniques, unacceptable as holotypes by today’s standards. The remainder of the series, which
he called ‘co-types’, were rarely labelled as such. His normal practice was to label very few, if any,
of these additional specimens, whether from the collection of the British Museum (Natural
History) or borrowed by him from elsewhere.
In the course of evaluating the type status of Arrow material for routine curation of the collec-
tion and for an increasing number of inquiries during the past few years, it was found that the
data quoted in his descriptions are rarely comprehensive. The localities given are often a précis
of the data labels of the series. Instances have been noted of published data, especially bionomic
notes, apparently from oral sources or contemporary correspondence, not appearing on the
labels. The name of the ‘collector’ may be the donor of the collection rather than the actual
collector. The number and sex of the specimens are usually omitted. These circumstances,
together with the fact that a considerable number of syntypes and paratypes must have been
given away or exchanged without record, make it impossible in many cases to be certain that the
whole of the series is accounted for. This paper, containing details of the type-material of 266
taxa, is the second of a series (cf. Bacchus, 1974) which will catalogue all the traceable type-
specimens of species described by G. J. Arrow. Of the total of about 1800 species described
Bull. Br. Mus. nat. Hist. (Ent.) 37 (3): 97-115 Issued 25 May 1978
97
by Arrow, approximately 760 Scarabaeidae and 650 species of 21 other families remain to be
catalogued.
Without access to the collection, museum registers of donations, various journals and lists
available only in the museum libraries, and some knowledge of the history of the collection and
of the vagaries of Arrow’s method, accurate and consistent collation of his type-series is virtually
impossible.
All specimens are in the British Museum (Natural History) except where otherwise stated.
The original genus of every species is given in parentheses after the specific name, and the current
valid name, when it differs from the original, is given at the end of the entry. The number and
sex of the original series are given in square brackets whenever possible. The data quoted are
not verbatim but have been rationalized for the sake of clarity, brevity and convenience to users.
Where possible the present name of the country of origin is used and all available data arranged
in a standard form. The museum registration numbers have been omitted as they are of no use
without the relevant registers.
The specimens labelled by Arrow as ‘types’ have, until now, all been regarded as holotypes
and all bore the round, red-bordered labels in use in this museum. Where lectotypes are desig-
nated they will, except where otherwise stated, be these specimens. Normally when both male and
female ‘types’ were so-labelled by Arrow the male is designated lectotype. In some of the genera
included in this catalogue it is not possible to sex accurately the specimens without dissection.
In most of these instances lectotypes have not been designated as it is considered that such dissec-
tion and subsequent lectotype designation are better left for specialist workers in the future.
In the few cases where it was considered expedient to designate lectotypes of unknown sex the
reasons for the decision are given in the text.
Every specimen catalogued in this paper will have a round label edged in red (holotype),
yellow (paratype), purple (lectotype) or blue (paralectotype or syntype) as appropriate and a
label with the name of the taxon written by the cataloguer and with his name printed below.
All cases where Arrow gave details of material in other collections were investigated and much
useful information received. Although in some instances the specimens could not be found (and
are noted as missing in the catalogue), it is probable that, partly because of Arrow’s habit of not
labelling many specimens and partly because of the considerable time such searches can take,
many of these specimens will eventually be found.
No information is available about material in the Colombo Museum.
Abbreviations
ANIC Australian National Insect Collection, Canberra.
BMNH _ British Museum (Natural History), London.
BPBM Bernice P. Bishop Museum, Honolulu.
FRI Forest Research Institute, Dehra Dun.
iP Institut fiir Pflanzenschutzforschung, Eberswalde.
IRSNB seInstitut Royal des Sciences Naturelles de Belgique, Brussels.
MCSN __ Museo Civico di Storia Naturale, Genoa.
MHNH Muséum National d’Histoire Naturelle, Paris.
MNHU_ Museum fiir Naturkunde der Humboldt Universitat, Berlin.
MZUSP Museu de Zoologia da Universidade de Sao Paulo, Sado Paulo.
SMT Staatliches Museum fiir Tierkunde, Dresden.
ZSI Zoological Survey of India, Calcutta.
Acknowledgements
I am indebted to Dr C. Costa, Dr R. Damoiseau, Mons. A. Descarpentries, Dr Swaraj Ghai,
Dr R. Hertel, Dr F. Hieke, Dr R. Krause, Mons. J. P. Lacroix, Dr H. Lyr, Dr N. C. Pant,
Dr R. Poggi, Dr P. Singh, Dr T. Sen Gupta, Dr E. Tortonese and Dr M. Uhlig for searching
their collections, providing data and lending specimens. I also thank Mr R. D. Pope for his
helpful advice and Miss V. Dick for preparing the index of species described by Gilbert Arrow
on which this catalogue is based.
98
Catalogue of the type-specimens described by G. J. Arrow
SCARABAEINAE
abreui (Onthophagus), 1931b : 239. LECTOTYPE 4, INpi<: Betul District, Dharakoh, 2600 ft, 25.iv.1926
(E. A. d’ Abreu); here designated.
Paralectotypes. INDIA: 1 3, same data as lectotype; 1 3, 19, same data as lectotype but 23.iv.1926;
1 g, Nagpur District, Pench Valley, Sylari, 1900 ft, 26.iv.1929 (E. A. d’Abreu); 1 9 Malabar (Doherty);
1 [sex unknown], Haldwani, Chakrata, ex dung, 4.v.1930 (B. M. Bhatia) (FRI, Dehra Dun).
acuticollis (Onthophagus), 1931b: 237. LECTOTYPE 3, INnpIA: Chikkaballapura (7. V. Campbell);
here designated.
Paralectotypes. INDIA: 1 9, Cape Comorin, 1888 (R. P. Castets).
[Specimen(s) from Madura (7. V. Campbell) missing.]
Current valid name: Onthophagus amphioxus Arrow (Arrow, 1933b : 421).
affinis (Arrow) var. capricornis (Liatongus), see capricornis.
affinis (Oniticellus), 1908 : 181. LECTOTYPE 3, BurMA: Tavoy; here designated.
Paralectotypes. INDIA: 1 3, Manipur. THAILAND: 2 ¢, 1 9, Renong (Doherty).
Current valid name: Liatongus affinis (Arrow) (Arrow, 19315 : 368).
agaricophilus (Onthophagus), 1931b : 333. LECTOTYPE 4, INDIA: Kumaon, W. Almora Div., vi.1917
(H. G. C[hampion]); here designated.
Paralectotypes. INDIA: 1 92, same data as lectotype but viii.1917; 1 3, same data as lectotype but
ix.1920; 1 9, Kumaon, C. Almora Div (H. G. C[hampion]); 1 2, Kumaon, Ranikhet Div, on Agaric,
vi.1921 (H. G. C[hampion]); 1 3, Ranikhet, vi-viii.1916 (H. G. C[hampion]); 1 [sex unknown],
Mussoorie, 600-700 ft, ix.1921 (Col. Harcourt) (FRI, Dehra Dun).
amphinasus (Onthophagus), 1931b: 195. LECTOTYPE 42, INpIA: Nilgiri Hills (H. L. Andrewes); here
designated.
Paralectotypes. INDIA: 1 3, 3 2, same data as lectotype; 1 3, same data as lectotype but 4000 ft,
vii.1908; 1 9, Nilgiri Hills (A. K. W. Downing); 1 [sex unknown], N. Kanara, Talewadi, nr Castle
Rock, 3.xi.1916 (S. Kemp) (ZSI, Calcutta).
andamanus (Paraphytus), 1931b : 419. Holotype (‘probably a 3’), ANDAMAN ISLANDs (Roepstorff ).
andrewesi (Onthophagus), 1931b : 324. LECTOTYPE 4, INpIA: Nilgiri Hills (A. K. W. Downing); here
designated.
Paralectotypes. INDIA: 3 3, 1 2, Nilgiri Hills (H. L. Andrewes); 4 3,1 9, Kanara; 3 3,2 9, Madras,
Anamalai Hills, 2400 ft, 6.v.1930 (J. C. M. Gardner).
angolensis (Coptorrhina), 1906 : 129. LECTOTYPE ¢, ANGOLA; here designated.
Paralectotypes. ANGOLA: 2 3.
[Arrow’s ‘9’ was the small male here designated as a paralectotype. The orange labels on all the
specimens refer to the register entry 1876-28 where the rest of the data can be found.]
angusticornis (Copris), 1933b : 427. Holotype 3, VIETNAM: Tonkin, Chapa, 25.iv.1918 (Jeanvoine).
antillarum (Onthophagus), 1903: 510. LECTOTYPE 3, St VINCENT: windward side (H. H. Smith);
here designated.
Paralectotypes. ST VINCENT: 2 3, 2 9 (H. H. Smith); 1 9°, South End (H. H. Smith). GRENADA:
1 9, windward side, Mount Gay Estate (H. H. Smith); 1 9, windward side, Balthazar (H. H. Smith);
1 9, no other data.
araneolus (Sisyphus), 1927a : 464 [original series: 2 ¢]. LECTOTYPE 4, INp1A: Nilgiri Hills, Gudalur,
v.1930 (E. E. Green); here designated.
Paralectotype. INDIA: 1 ¢ same data as lectotype.
arboreus (Onthophagus), 1931b : 225. LECTOTYPE 4, INp1A: Bihar, Pusa, on grass, 28.vii.1922 (Ram[rest
illegible]); here designated. uf
Paralectotypes. INDIA: 1 9, Bihar, Pusa, 14.ix.1912 (R.M.P.); 1 3, Dehra Dun, New Forest, 27.11.1929,
on sand (J. C. M. Gardner).
[Specimens from INDIA: Sagar, xi; Pusa, viii, missing. ]
argentinum (Pedarium), 1913a : 459. Syntypes, ARGENTINA: 4 ex., Rio Salado, Gran Chaco (E. Wagner);
3 ex., Rio Salado, Icano (E. Wagner).
assamensis (Panelus), 1907c : 417 [original series: 3]. Syntypes, INDIA: 2 ex., Assam, Patkai Mts (Doherty);
1 ex., Assam, Sudiya (Doherty).
99
aviculus (Onthophagus), 1931b: 179. LECTOTYPE 4, Inp1A: Bengal, Darjeeling, Singla, 1500 ft, vi;
here designated.
Paralectotypes. INDIA: 4 3, 4 9 (BMNH, London), 1 [sex unknown] (ZSI, Calcutta), same data as
lectotype; 1 2, Darjeeling, Tukvar (E. A. Moller). BURMA: 3 3, 2 2, Putao Dist., Sumparbum, vi-vii
(Mrs B. Fischer).
[Specimen(s) Darjeeling, Tinta Valley, 700 ft, ix (J. C. M. Gardner) missing.]
Current valid name: Onthophagus anguliceps Boucomont (Arrow, 19336 : 421).
avocetta (Onthophagus), 1933b : 421 [proposed by Arrow as replacement name for Onthophagus anguliceps
Boucomont sensu Arrow (19315: 178)]. LECTOTYPE 3, InpIA: Assam, Garo Hills above Tura,
3500 ft, 15.vii-30.viii.1917 (S. Kemp); here designated.
Paralectotypes. INDIA: 3 9, same data as lectotype; 1 [sex unknown], same data as lectotype but
ix.1917 (Mrs Kemp). BURMA: 1 3, Karen Hills, Cheba, 900-1100 m, v-xii.1888 (L. Fea).
beesoni (Onthophagus), 19316 : 319. LECTOTYPE 4, INDIA: Kumaon, Haldwani Dist. (H. G. Champion);
here designated.
Paralectotypes. INDIA: 2 ¢ (BMNH, London); 1 [sex unknown] (FRI, Dehra Dun);2 ¢ Jhaira Range,
Dehra Dun, 6.xi.1924 (C. F. C. Beeson); 1 3, Jhaira, Dehra Dun, ex dung, 14.ii.1929 (J. C. M. Gardner).
[Specimen(s) from INpDIA: Bengal, Buxar Duars, v (D. Nowrojee) missing.]
biharensis, Gymnopleurus, 1931b : 62. LECTOTYPE 4, INDIA: Chapra (Mackenzie); here designated.
Paralectotypes. INDIA: 1 3, 2 2, same data as lectotype; 1 3, Bengal, Chapra.
[Specimens in the Pusa collection (now Indian Agricultural Research Institute, New Delhi) missing.]
bisectus, (Onthophagus), 1931b : 201. LECTOTYPE 4, INp1A: S. India; here designated.
Paralectotypes. INDIA: 1 2, same data as lectotype; 1 3, Malabar; 2 ¢, 3 9, Nilgiri Hills (A. K. W.
Downing).
[Specimen(s) from INDIA: Snowdon Peak, 8000 ft, ix (Y. R. Rao) missing.]
bombayensis (Gymnopleurus), 1931b : 65 [original series: 6]. LECTOTYPE 4, no locality label, presented
by Walter Elliot Esq.; here designated. ;
Paralectotypes. 1 3, same data as lectotype; 1 3, ‘Quetta Museum’. PAKISTAN: 1 9, Baluchistan,
500-800 ft, 26.vii—6.viii.1929 (J. W. Evans). INDIA: 1 9, Belgaum, Bombay; 1 9, Ind. Cent.
brachialis (Uroxys), 1933a : 393. Holotype 3, CoLumBIA: Pichinde, 5000 ft, xi.1891 (W. Rosenberg).
brevicollis (Onthophagus), 1907c : 422. LECTOTYPE 4, INpIA: Nilgiri Hills; here designated.
Paralectotypes. INDIA: 1 3, 1 2, same data as lectotype; 1 3, 1 9, same data as lectotype but (G. F.
Hampson); 2 3, 2 9, same data as lectotype but (H. L. Andrewes); 1 3, Belgaum, Bombay; 1 2,
Malabar; 1 9, South India.
bronzeus (Onthophagus), 1907c : 429. LECTOTYPE 32, Inp1A: S. India; here designated.
Paralectotypes. INDIA: 1 9, Nilgiri Hills (G. F. Hampson); 1 9°, S. India, Kanara; 1 2, Malabar.
brutus (Onthophagus), 1931b : 215. LECTOTYPE 4, INp1A: Bengal, Sarda (F. W. C[hampion)).
Paralectotype. BURMA: 1 9, Carin Cheba, 900-1100 m, v—xii.1888 (L. Fea).
bucki (Zonocopris), 1932 : 223. Syntypes, BRAZIL: 1 ex., Porto Alegre, Rio Grande do Sul, x.1931 (P. Buck);
1 ex., as previous but iv.1931; 6, as previous but dates unknown (MZUSP, Sao Paulo).
[Although Arrow does not give the precise number of specimens seen by him he does note that there
were many examples and that 12 specimens were found inside one snail shell.]
buffalo (Onthophagus), 1907c : 437. LECTOTYPE 4, JAVA: here designated.
Paralectotypes. 1 3, 4 2, same data as lectotype. BoRNEO: 1 9, Labuan; 1 3, 1 2, Borneo [rest of
label illegible].
bufo (Onthophagus), 1907c : 420. LECTOTYPE 4, PAKISTAN: Bannu (Dr Pennell); here designated.
Paralectotypes. PAKISTAN: 6 3, 6 9, same data as lectotype; 1 3,2 9, same data as lectotype but no
collector. INDIA: 1 9°, Gwalior.
bufulus (Onthophagus), 1941: 51. Holotype 3, New Guinea: Papua, Mafulu, 4000 ft, i.1934 (L. E.
Cheesman).
calcaratus (Pinotus), 1913a: 457. Holotype 3, BraziL: Rio Madeira, Madeira-Mamore R[ail] R[oad]
Co., Camp 41 (Mann & Baker).
callosipes (Sisyphus), 1909a: 517. LECTOTYPE 3, MozaAmsiquE: Nyassa (Simons); here designated
[Arrow gives this locality as British Central Africa, Nyassaland (Malawi), apparently not realizing
that part of L. Nyasa is, in fact, in Mozambique].
100
Paralectotypes. MOZAMBIQUE: 1 3, same data as lectotype; 1 3g, 1 9, ‘Nyassa’ [it is not possible to
assign these specimens to the Mozambique or the Malawi part of ‘Nyassa’]. RHODESIA: 1 9, Mashona-
land, Mt Chirinda, xi.1901 (G. A. K. Marshall). TANZANIA: 1 3, Massai (sic). ZAIRE: 1 3, 1 2, 150-
200 mls W. of Kambove, 3500-4500 ft, 18.x.1907.
cancer (Mnematium), 1919 : 434. Holotype 3, no locality label, Nevinson Collection.
capricornis (Liatongus affinis (Arrow) var.), 19316 : 368 [original series: 4 3, unspecified number of 9].
LECTOTYPE 3, BurMa: Pegu, Palon, viii—x.1887 (L. Fea); here designated.
Paralectotype. BURMA: 1 3, Tharawaddy (G. Q. Corbett).
Current valid name: Liatongus capricornis Arrow (Janssens, 1953, Explor. Parc natn. Upemba Miss.
G. F. de Witte 11 : 96).
caucanus (Uroxys), 1933a : 394. LECTOTYPE 3, CoLomsiA: Cauca; here designated.
Paralectotypes. COLOMBIA: 1 3, 2 9, same data as lectotype.
cheesmanae (Onthophagus), 1941 : 52. LECTOTYPE 3, New Guinea: Papua, Kokoda, 1300 ft, ix.1933
(L. E. Cheesman); here designated.
chrysurus (Onthophagus), 1931b: 211. LECTOTYPE 3, INpIA: Chachar, Haflong, 15.v.1925 (G. D.
Bhasin); here designated.
[This specimen has been designated lectotype because it is uncertain that it is unique.]
cingalense (Phacocosma) 1931b : 357. Holotype [sex unknown], Sri LANKA: Ohiya, 6500 ft, 9.iv.1928.
circulifer (Onthophagus), 1931b : 274. LECTOTYPE 4, Inp1A: Dehra Dun, 7.vii.1925 (C. F. C. Beeson);
here designated.
Paralectotypes. INDIA: 1 2, Darj[eeling], Soom, 400-500 ft, 8.vii.1914; 1 3, Bengal, Darjeeling,
Singla, 1500 ft, vi; 1 9, Siwalik Hills, Mohand Rau, 23.ix.1921 (Dr Cameron); 1 9, Jummu Prov,
Chenab V, Ramban, 2600 ft, 8.v.1928 (C. F. C. Beeson); 4 3,2 9, Sikkim, Pedong, 6000 ft, 28.iii.1924
(Maj. R. W. G. Hingston). BuRMA: 1 2? (BMNH, London); ? number and sex (FRI, Dehra Dun);
Chittagong H[ill] T[racts], Sitaphar R, 5-10.v.1925 (C. F. C. Beeson).
In FRI, Dehra Dun, there are also specimens of unknown number and sex from INDIA: Assam,
Shillong (C. F. C. Beeson); Jhaira Range, Dehra Dun, 6.xi.1925 (C. F. C. Beeson).
[Specimens from INp1A: Khinsi Hills, Nagpur, 1500 ft, ix (E. A. d’Abreu); Darjeeling District, Lord
Carmichael’s collector, missing.]
coeruleicollis (Onthophagus), 1907c : 430. LECTOTYPE 2, INDIA: Dhawar; here designated.
Paralectotypes. INDIA: 1 g, 1 2, Belgaum, Bombay; 1 9, India.
coorgensis (Onthophagus), 1931b : 202. LECTOTYPE 4, INnpIA: Nilgiri Hills (H. L. Andrewes); here
designated.
Paralectotypes. INDIA: 5 3, 8 9, same data as lectotype.
[Specimen(s) from INDIA: Malabar Dist., Sappal, Palghat, 1700 ft, v (J. C. M. Gardner) missing.]
cristatum (Trichillum), 1931a : 610 [‘taken in considerable numbers by Dr. O’haus’]. Syntypes, ECUADOR:
1 ex., Mahader Stadt, Loja, 2200 m (F. O’ Haus); 1 ex., Loja, 1.x.1905 (F. O’ Haus); 1 ex., Piscobamba
(F. Witt).
cryptops (Pedarium), 1913a : 458. Syntypes, BRAZIL: 2 ex., Goyaz Prov, Jatahy; 1 ex., Natal (W. M.
Mann).
[One specimen, BRAZIL: Natal (W. M. Mann) missing.]
capreiceps (Onthophagus), 1907c : 427. LECTOTYPE 4, Inp1a: Sikkim, Teesta Valley, Tungu, 1300-
1400 ft, 1-15.vii.1903; here designated.
Paralectotypes. INDIA: 1 3, 3 2, same data as lectotype; 1 3, 2 2, Khamba Jong, 1500-1600 ft,
15-30.vii.1903.
digitatus (Onthophagus), 1931b : 177. LECTOTYPE 4, INp1A: ‘Ind. bor.’; here designated.
Paralectotypes. INDIA: 1 9, same data as lectotype; 1 3, Sikkim (MNHN, Paris).
draco (Copris), 1906 : 127. LECTOTYPE 3, ANGOLA: Bihe; here designated.
Paralectotypes. ANGOLA: 2 9, same data as lectotype.
delicatus (Copris), 1931b:117 [original series: 2]. LECTOTYPE 6d, INDIA: Mishmi Hills, 28°5’ N,
96°15’ E, 2000 ft, 12.iii.1928; here designated.
Paralectotype. INDIA: 1 2, Bengal, Darjeeling, Lopchu, 5000 ft, 24.ix.1929 (J. C. M. Gardner).
dynastoides (Onthophagus), 1931b : 182. Holotype 3, INDIA: ‘Ind. bor.’.
101
egregius (Onthophagus), 1907c : 438. LECTOTYPE 3, Borneo: Pontianak, 1893; here designated.
Paralectotypes. BORNEO: 1 3, 1 2 [not ¢] (BMNH, London), 1 9 [not 3] (MNHN, Paris), same data
as lectotype but 1898; 1 2 [not 3] (BMNH, London), 1 2 [not 3] (MNHN, Paris), same data as
lectotype but 1900; 1 9 (BMNH, London), 1 2? (MNHN, Paris) [both described by Arrow as the female
of egregius], same data as lectotype but 1898.
[The specimens described by Arrow as the female of Onthophagus egregius are Onthophagus aurifex
Harold (Balthasar, 1963, Monographie der Scarabaeidae und Aphodiidae der Palaearktischen und
Orientalischen Region 2 : 340).]
elongatus (Gyronotus), 1933b : 429. LECTOTYPE 3, SourH Arrica: Natal, N. Zululand; here desig-
nated.
Paralectotypes. SoUTH AFRICA: 1 3, 2 92, same data as lectotype.
Current valid name: Gyronotus dispar Felsche (Janssens, 1938, Expl. Parc natn. Albert Miss. G. F.
de Witte 21: 16).
ephippiatus (Deronitis), 1933b : 422. Holotype 3, UGANDA: Kasinga Channel, Katunguru, 1931 (Dr E. B.
Worthington).
ephippioderus (Onthophagus), 1907c : 425. LECTOTYPE <2, INpIA: Bangalore; here designated.
Paralectotypes. INDIA: 5 3, 69, same data as lectotype; 5 3, 3 9, Mysore; 2 3, Nilgiri Hills (G. F.
Hampson); 4 3,9 9, Nilgiri Hills; 1 3, 1 9, Belgaum, Bombay; 1 3,1 9, Belgaum; 2 3,3 92, Kanara.
excavatus (Onitis), 1931b : 390. LECTOTYPE 3, BurMa: Tenasserim; here designated.
exquisitus (Onthophagus), 1931b : 225. LECTOTYPE 2, INpIA: Dehra Dun, 14.vii.1922 (S. N. Chatterjee);
here designated.
Paralectotype. INDIA: Bengal, Darjeeling, Singla, vi.
felix (Canthon), 1913a: 456. LECTOTYPE <3, BrRAziL: Santarem; here designated.
Paralectotypes. BRAZIL: 1 9, same data as lectotype; 1 ¢, 3 9, Pernambuco (A. Fry); 1 3, Para;
1 3, Ceara, Baturite Mts (W. M. Mann).
[Specimens from BRAZIL: Rio de Janeiro; Ceara, Maranguape Mts missing. ]
felix (Onthophagus), 1931b : 257. Holotype 3, INDIA: Chota Nagpur, Nowatoli, x.1897 (R. P. Cardon).
Paratypes. INDIA: 4 2, same data as holotype.
fossor (Onthophagus), 1931b: 300. Holotype 3, INDIA: Sikkim, Tsuntang, 6000 ft, 24.iv.1924 (Maj.
R. W. G. Hingston).
Paratypes. INDIA: 1 ¢ 3 9, same data as holotype.
frugivorus (Onthophagus), 1931b : 336. LECTOTYPE 4, INpIA: Assam, Garo Hills above Tura, 3500-
3900 ft, 15.vii—30.viii.1917 (S. Kemp); here designated.
Paralectotypes. INDIA: 1 9, same data as lectotype; 4 [sex unknown], same data as lectotype but
15-25.vii.1917, in rotten fruit (ZSI, Calcutta); 1 2, same data as lectotype but 3900 ft, 15.vii—-15.viii.1917;
2 3, 1 9, as previous but in rotten fruit.
fulgens (Pedarius), 1913a : 458 [original series: 2]. Syntypes, ARGENTINA: 2 ex., Gran Chaco, Rio Salado
(E. Wagner).
furcicollis (Onthophagus), 1931b : 276. LECTOTYPE <2, INpIA: Gopaldhara, Darjeeling, 3440-4720 ft,
19.ix.1916, on ground (H. Stevens); here designated.
Paralectotype. INDIA: 1 ex., Dehra Dun, Nun Nadi, ex carrion, 17.vii.1921 (Dr Cameron).
gallinus (Caccobius), 1907c : 424. LECTOTYPE 2, INp1A: Nilgiri Hills (H. L. Andrewes); here designated.
Paralectotypes. INDIA: 16 3, 17 9, same data as lectotype; 1 9, same data as lectotype but 600 ft;
7 3,10 9, Nilgiri Hills; 1 2, Belgaum; 2 9°, Belgaum, Bombay.
gazanus (Onitis), 1909a: 519. LECTOTYPE 3, RuopesiA: Mashonaland, Chirinda Forest, x.1908
(G. A. K. Marshall); here designated. ;
Paralectotypes. RHODESIA: 7 3, 4 9, same data as lectotype.
[By modern definition, Chirinda Forest is not in Mashonaland. It is 110 miles south of Umtali.]
gazanus (Sisyphus), 1909a : 518. LECTOTYPE 3, RHopeEsIA: Mashonaland, Chirinda, x.1905 (G. A. K.
Marshall); here designated.
Paralectotypes. RHODESIA: 1 3, Mashonaland, Mt Chirinda, xi-xii.1901 (G. A. K. Marshall); 1 3,
as previous but xii.1905; 1 9, Gazaland, Chirinda, xii.1901 (G. A. K. Marshall). MOZAMBIQUE: 2 3,
Lower Buzi River, Chibababa, xii.1906 (C. F. M. Swynnerton).
[The female paralectotype was re-determined as Sisyphus ocellatus Reiche by E. Haaf. There is some
evidence that the words ‘co-type’ have been cut from the bottom of Arrow’s determination label.]
102
geminatus (Pinotus), 1913a : 457 [original series: 2]. LECTOTYPE 3, Brazi_: Natal (W. M. Mann);
here designated.
gilleti (Pseudopedaria), 1933b : 424. LECTOTYPE 3, RuopesiA: Kashitu, N. of Broken Hill, ii.1915
(H. C. Dollman); here designated.
Paralectotypes. RHODESIA: 2 2, same data as lectotype but xii.1914; 4 9, same data as lectotype but
i.1915; 1 9, same data as lectotype but iv.1915. TANZANIA: 2 9.
gilleti (Synapsis), 1931b:83. LECTOTYPE 9, INpIA: Bengal, Darjeeling, Singla, 1500 ft, vi; here
designated.
Paralectotype. INDIA: 1 ¢ (IRSNB, Brussels).
gladiator (Copris), 1933b : 425. Holotype 3, BURMA: Adung Valley, 6000 ft, 11.v.1931 (Lord Cranbrook).
gladiator (Onthophagus), 1907c : 418 [original series: 2]. LECTOTYPE 3, INpIA: here designated.
Paralectotype. INDIA: 1 3, same data as lectotype.
gladiator (Sisyphus), 1927a : 462. LECTOTYPE 3, CAMEROUN: Ngpwdar; here designated.
Paralectotypes. CAMEROUN: 4 3, 4 9, same data as lectotype; 1 9, Tazada.
[Specimen(s) from CAMEROUN: Betare missing.]
globosus (Parachorius), 1931b : 360. Holotype 9 ?, INDIA: Darjeeling, 5000 ft, 14.iii.1924 (Maj. R. W. G.
Hingston).
gorgon (Uroxys), 1933a: 397 [original series: ‘more than a dozen’]. LECTOTYPE 6, CoLomsia:
Gorgona Island, in hair of three-toed sloth, 3.vii.1924; here designated.
Paralectotypes. COLOMBIA: 2 3, 6 2, same data as lectotype; 1 2, same data as lectotype but (H. J. K.).
gratus (Onthophagus), 1931b : 212. LECTOTYPE 4, Inp1a: ‘Ind. bor.’; here designated.
Paralectotypes. INDIA: 1 2 (BMNH, London), 1 [sex unknown], (ZSI, Calcutta), Ranchi (W. H.
Irvine); 1 3, Ranchi.
griseosetosus (Onthophagus), 19316: 192. LECTOTYPE 4g, INpDIA: South Mysore (H. L. Andrewes);
here designated.
Paralectotypes. INDIA: 1 g, 2 9, same data as lectotype; 2 3, Nilgiri Hills (H. L. Andrewes); 1 2,
Kanara; 1 3, C[entral] P[rovince], Raigarh R., vii.1927 (C. F. C. Beeson).
[Specimen(s) from INDIA: Dehra Dun, vi, missing.]
gulo (Onthophagus), 1931b : 164. LECTOTYPE 3, INDIA: Bombay; here designated.
Paralectotypes. INDIA: 2 2, same data as lectotype; 1 9, Nagpur (E. A. D’Abreu); 1 2, Buldana,
Purna R, Sagoda, 21.viii.1929 (N. C. Chatterjee); 1 9, as previous but 7.ix.1929; 1 9, as previous but
8.1ii.1930; 6 [sex unknown], as previous but 26.vi.1929 [FRI, Dehra Dun]; 1 [sex unknown], as previous
but 4-13.x.1921 [FRI, Dehra Dun].
The dates of the paralectotypes in FRI, Dehra Dun, were not listed by Arrow although they are
labelled “Cotypes’ and are from the same series as the specimens from the same locality in the BMNH,
London. The dates quoted by Arrow are only those of the BMNH specimens and the omission of the
data of the FRI specimens is considered to be an accident caused by Arrow’s habit of reducing data to
an absolute minimum.
hamaticeps (Onthophagus), 1931b : 308. LECTOTYPE 32, INp1A: Siwaliks [hills], Dehra Dun, 24.ix.1922
(Dr Cameron); here designated.
Paralectotype. INDIA: 1 3, Jhaira Range, Dehra Dun, 6.xi.1924 (C. F. C. Beeson).
hindu (Onthophagus), 19316 : 289. LECTOTYPE 3, INp1A: ‘India Orient.’; here designated.
Paralectotypes. INDIA: 5 9, same data as lectotype. PAKISTAN: 1 3, Bannu (Dr. Pennell).
[Specimens from INDIA: Nagpur, 1000 ft, viii & ix (E. A. D’ Abreu) and Bellary Dist., Yemmiganur,
viii, missing. ]
hindu (Paraphytus), 19136 : 193. Holotype 3, INDIA: Khasia Hills, vii.1894.
Paratypes. INDIA: 2 3, same data as holotype; 2 9, Upper Rotung, under bark, 6.i.1912 (Kemp).
[1 3, same data as previous, missing.]
hingstoni (Onthophagus), 1931b:278. LECTOTYPE 4, INpIA: Sikkim, Lachen, 9000 ft, 26.iv.1924
(Maj. R. W. G. Hingston); here designated.
Paralectotypes. INDIA: 1 g, same data as lectotype but 8000 ft, 24.iv.1924; 2 2, same data as lecto-
type but 10 000 ft, 25.iv.1924.
[Specimen(s) INDIA: Sikkim, Tsuntang, 7000 ft (Maj. R. W. G. Hingston) missing.]
histeroides (Uroxys), 1933a : 392. LECTOTYPE 3, PARAGUAY: here designated. Paralectotypes. PARA-
GUAY: 2 @.
103
hookeri (Parachorius), 1931b : 359. Holotype 2 ?, INDIA: N. India.
humeralis (Cassolus), 1907c : 416. LECTOTYPE 4, INpIA: Assam, Sudiya (Doherty); here designated.
Paralectotypes. INDIA: 1 3, 2 2, Patkai Mts (Doherty); 2 9, Valley (Doherty).
[One paralectotype, INDIA: Patkai Mts (Doherty) lost, empty mount in BMNH, London.]
hystrix (Trichillum), 1931a: 609. Syntypes, ARGENTINA: 2 ex., Santa Fé, Rio San Janvier, Estancia la
Noria, xii.1911 (G. E. Bryant); 1 ex., as previous but 14.xii.1911; 1 ex., as previous but 20.xii.1911;
1 ex., as previous but 23.xii.1911; 1 ex., as previous but 24.xii.1911; 1 ex., as previous but 27.xii.1911;
1 ex., as previous but 3.i.1912.
indicus (Oniticellus), 1908 : 180. LECTOTYPE 4, Inp1A: Nilgiri Hills; here designated.
Paralectotypes. INDIA: 3 3, 1 9, same data as lectotype; 1 g, S. India; 1 g, 1 9, Anamalais; 1 9,
Anamalai Hills; 2 3, 3 9, Malabar.
Current valid name: Chironitis arrowi Janssens (Janssens, 1937, Mém. Mus. r. Hist. nat. Belg. (2)
11 : 159).
inermis (Caccobius), 1931b : 147. LECTOTYPE 4, INpDIA: Fyzabad (R. W. G. Hingston); here designated .
Paralectotypes. INDIA: 1 3, Nagpur (E. A. D’ Abreu); 2 3, W. Almora, Kumaon (H. G. C[hampion]);
1 9, Calcutta; 1 9, Sikkim, Pedong, 6000 ft, 28.iii.1924 (R. W. G. Hingston). SR1 LANKA: 1 3, Wirawila,
28.vii.1921.
iyengari (Onthophagus), 1931b : 323. Holotype 3, INDIA: Bengal, Rungpur, xi.1924 (M. O. T. Iyengar);
here designated.
Paratypes. INDIA: 1 3, 2 2, same data as holotype.
jucundus (Onthophagus), 1931b : 188. LECTOTYPE 3, INDIA: Assam, Manipur (Doherty); here desig-
nated.
Paralectotype. BURMA: 1 9, Momeit [not Momeik] (Doherty).
kanarensis (Onthophagus), 1931b : 191. LECTOTYPE 4, INDIA: N. Kanara Dist., Talewadi, nr Castle
Rock, 3-10.x.1916 (S. Kemp); here designated.
Paralectotype. INDIA: 1 9, Kanara. ;
kumaonensis (Onthophagus), 1931b:272. LECTOTYPE 3, INDIA: Kumaon, Nainital Div. (H. G.
C[hampion]); here designated.
Paralectotypes. INDIA: 1 3, same data as lectotype; 1 2, W. Almora, v—viii.1916 (H. G. C[hampion));
1 9, Simla Hills, Theog, 7600 ft, 10.ix.1921 (S. N. Chatterjee); 1 3, Mussoorie, vii.1926 (J. C. M.
Gardner); 1 9, as previous but vi.1930; 1 2, Mussoorie, 6500 ft, 6.vii.1920 (S. N. Chatterjee); 1 [sex
unknown], Mussoorie, 6000-7000 ft, xi.1921 (Col. Harcourt) (FRI, Dehra Dun).
laborans (Onthophagus), 1931b : 240. LECTOTYPE 32, INpIA: Nasik, 31.vii.1911 (NV. B. Kinnear); here
designated.
Paralectotypes. INDIA: 2 3, 4 9, Chikalda Berars, 3664 ft, 2.xii.1913 (N. B. Kinnear); | er tae he A
Dharwar, x.1909 (H. Swale); 2 9, as previous but xii.1909; 2 9, as previous but 17.i.1910.
laetum (Phacosoma), 1931b : 356. Holotype [sex unknown], INp1A: Nilgiri Hills (H. L. Andrewes).
Paratypes. INDIA: 4 [sex unknown], same data as holotype.
laevis (Gymnopleurus), 1900: 23. Holotype 3, SoMALI REPUBLIC: ‘Somaliland’, 16.iv—7.viii.1895 or
5.vi-29.x.1897 (C. V. A. Peel).
Current valid name: Garreta nitens (Olivier) (Janssens, 1940, Mem. Mus. r. Hist. nat. Belg. (2) 16 : 32).
laevis Harold var. stevensi (Onthophagus), see stevensi.
laoticus (Sisyphus), 1927a : 463. LECTOTYPE 3, INpo-Cuina: Haut Mekong, Vieng Vien, 24.v.1918
(R. V. de Salvaza); here designated. ;
Paralectotypes. INDo-CHINA: 1 3, 2 9, same data as lectotype; 1 9, Haut Mekong, Vien Vai,
23.v.1918 (R. V. de Salvaza).
lapillus (Onthophagus), 1931b : 267. LECTOTYPE 4, INpIA, Sikkim, Pedong, 6000 ft, 28.vii.1924 (Maj.
R. W. G. Hingston); here designated.
Paralectotypes. INDIA: 2 2, same data as lectotype; 1 3g, 1 9, Rungbong Valley, Gopadhara (H.
Stevens); 1 9, Dehra Dun, 18.viii.1915; 1 [sex unknown], Dehra Dun, vi.1918 [FRI, Dehra Dun];
1 [sex unknown], Dehra Dun, under bark 13.viii.1927 (S. Badahur) [FRI, Dehra Dun]; 1 3, Mussoorie,
vii.1926 (J. C. M. Gardner); 1 [sex unknown], Mussoorie, Mackinnon Park, under leaves, 29.viii.1929
(C. F. C. Beeson) [FRI, Dehra Dun]; 1 [sex unknown], Kumaon, Shamkhet, 15.v.1912 [FRI, Dehra
Dun]; 1 2, Punjab, Simla Hills, Dagshai, 6000 ft, 13.viii.1921 (O. H. Walters); 1 9, N. India.
104
laratinus (Onthophagus), 1916 : 494 [original series: 3 3, 2 9]. Holotype 3, LARAT I. (Indonesia): (F.
Muir); here designated.
Paratypes. LARAT I. (Indonesia): 1 3, 1 2, same data as lectotype.
latenasutus (Onthophagus), 1941 : 50. LECTOTYPE 3, NEw GuineA: Papua, Kokoda, 1200 ft, ix.1933
(L. E. Cheesman); here designated.
Paralectotypes. NEw GuINEA: 7 3, 7 9, same data as lectotype; 1 3, D[utch] N[ew] Guinea
(Sattelburg).
latus (Uroxys), 1933a : 396. Holotype 3, BRAZIL: Petropolis, ii.1857 (H. Clark).
lojanus (Uroxys), 1933a: 395. LECTOTYPE 3, Ecuapor: Loja Pucara, 8.viii.1905 (F. Ohaus); here
designated.
Paralectotypes. ECUADOR: 1 2, same data as lectotype but 9.viii.1905; 1 9, Loja Clavario, 5.viii.1905
(F. Ohaus); 1 3, as previous but 1.ix.1905; 3 3, 3 2, Loja (F. Ohaus) (IP, Eberswalde).
madoqua (Onthophagus), 1931c : 258. Holotype 3, INDIA: Nilgiri Hills (H. L. Andrews).
Paratypes. INDIA: 2 3, 2 2, same data as holotype.
manipurensis (Onthophagus), 1907b : 426. LECTOTYPE 4, INDIA: Manipur (Doherty); here designated.
Paralectotypes. INDIA: 2 3, 2 2, same data as lectotype.
manni (Canthidium), 1913a : 460. LECTOTYPE 3, BraAziL: Ceara (W. M. Mann); here designated.
Paralectotype. BRAZIL: 1 9, same data as lectotype.
medicus (Sisyphus), 1931b : 70. Holotype 3 ?, INDIA: Madras, Anamalai Hills, 2400 ft, 10.v.1930 (J. C. M.
Gardner).
metalliceps (Onthophagus), 1931b : 218. LECTOTYPE 4, InpIA: Kanara; here designated.
Paralectotype. INDIA: 1 9, same data as lectotype.
minotaurus (Onthophagus), 1941 : 53 [original series: 1 g, 1 9]. Holotype 3, NEw Guinea: Papua,
Kokoda, 1200 ft, x.1933 (L. E. Cheesman).
Paratype. NEw GUINEA: 1 9, D[utch] New Guinea, Sattelburg.
mirandus (Onthophagus), 1931b : 290. LECTOTYPE 3, INDIA: W. Almora, Kumaon (H. G. C[hampion));
here designated.
Paralectotypes. INDIA: 6 3, 2 2, same data as lectotype; 1 3, 2 9, Simla, vi. [not vii] 1909 (H.
Chippendale).
modestus (Oniticellus), 1908 : 182. LECTOTYPE 4, INp1A: Calicut; here designated.
Paralectotypes. INDIA: 1 2 [head and pronotum missing], 7 2, same data as lectotype; 1 2 Belgaum;
1 3,1 9, Belgaum, Bombay; 2 3, 2 2, Malabar.
morio (Sisyphus), 1909b : 93. LECTOTYPE 3, Cuina: Ai San, 30 mls W. of Chefoo (M. P. Anderson);
here designated.
Paralectotypes. CHINA: 3 9, same data as lectotype; 1 9, Shan-hai-Kwan, in mountains, i.1906
(F. M. Thomson).
Current valid name; Sisyphus schaefferi L. (Haaf, 1955, Ent. Arb. Mus. Georg. Frey 6 : 344).
myrmecophilus (Onthophagus), 1907c : 431. LECTOTYPE 4, INp1A: Nilgiri Hills (H. L. Andrewes); here
designated.
Paralectotypes. INDIA: 4 3, 6 9, same data as lectotype; 1 g, same data as lectotype but ‘taken close
to nest of Phidole indica Mayr whilst I was digging out the nest. Probably lives with the ants’; 6 3,
2 2, Nilgiri Hills, Barwood Estate, 3500 ft, found with ants in decayed Ficus trunk living in galleries
of ants (Phidolegiton sp., probably diversus (Gerdon)), x.1906 (H. L. Andrewes).
nagpurensis (Onthophagus), 1931b : 342. Holotype 3, INDIA: Chota Nagpur, x.1897.
Paratypes. INDIA: 2 3g, 3 9°, same data as holotype.
nasalis (Onthophagus), 1931b : 288. Holotype 3, INDIA: W. Almora, Kumaon (H. G. C[hampion]).
necrophagus (Onthophagus), 1931b : 196. LECTOTYPE 4, INDIA: Dehra Dun, Nun Nadi, ex carrion,
17.vii.1921 (Dr Cameron); here designated.
Paralectotypes. INDIA: 1 3, 1 2, Dehra Dun, Jhajra, Sal leaves, 27-29.vi.1912 (C. N. Chatterjee).
nitidicollis (Oniticellus), 1908 : 179. Holotype 3, SARDINIA.
oculatus (Onthophagus), 1931b : 168. LECTOTYPE 4, Sri LANKA: Nilaveli, ix.1914 (C. F. S. Baker);
here designated.
Paralectotypes. Sk1 LANKA: 1 3, 1 2, same data as lectotype. INDIA: 1 2, Malabar; 1 92, Pondicherry.
105
ohausi (Trichillum), 1931a: 610 [original series: ‘considerable numbers’]. Syntypes, EcuADoR: 1 ex.,
Loja Punzara, 7.viii.1905 (F. Ohaus); 1 ex., Loja Calvario, 4.viii.1905 (F. Ohaus); 1 ex., Loja (F. Ohaus);
2 ex., (F. Ohaus).
onthochromus (Onthophagus), 1913a: 460. LECTOTYPE 3, BraziL: Rio Madeira, Madeira-Mamore
R[ail] R[oad] Co. Camp 41 (Mann & Baker); here designated.
Paralectotype. BRAZIL: 1 2, same data as lectotype.
orissanus (Onthophagus), 1931b : 257. LECTOTYPE 4, INnp1A: ‘E. Ind.’; here designated.
Paralectotypes. INDIA: 2 9, same data as lectotype; 1 3, Orissa, Barkul, 0—-1000 ft, 1-3.viii.1914
(Gravely).
paliceps (Onthophagus), 1931b : 287 [original series: 2]. Holotype 3, INDIA: ‘Ind. bor’.
[The paratype was probably from the collection at FRI, Dehra Dun.]
pantherinus (Caccobius), 1931b : 154. LECTOTYPE 3, PAKISTAN: Karachi; here designated.
Paralectotype. INDIA: 1 9, Punjab, Multon Div, Gazighat, ex cattle dung, 15.ii.1928 (B. M. Bhatia).
paranense (Pedaridium), 1932 : 224 [original series: 4]. Syntypes, BRAZIL: 4 ex., Parana, Castro.
peninsularis (Cassolus), 1907c : 437. LECTOTYPE 3, West MALAysIA: Perak (Doherty); here designated.
Paralectotypes. WEsT MALAYsIA: 2 9, same data as lectotype; 1 3, Penang (Lamb).
platalea (Onthophagus), 1941 : 49. Holotype 3, NEw GuINnEA: Japen I., R. Manai-Undei, 500 ft, x.1938
(L. E. Cheesman).
porcus (Onthophagus), 1931b : 325. LECTOTYPE 3, BANGLADESH: Chittagong H[ill] T[racts], Sitaphar
R., 5—10.v.1925 (C. F. C. Beeson); here designated.
Paralectotypes. BANGLADESH: 2 ¢ (BMNH, London), 2 ¢ (FRI, Dehra Dun), same data as lectotype.
[The female specimen described by Arrow is missing.]
prehensilis (Macropocopris), 1920 : 436. Holotype 3, AUSTRALIA: N. Queensland, Kuranda, 19.xi.1909
(G. E. Bryant).
Current valid name: Onthophagus prehensilis (Arrow) (Matthews, 1972, Aust. J. Zool. Suppl.
24 : 42).
productus (Onthophagus), 1907c : 423. LECTOTYPE 4, INp1A: (Capt. Boys); here designated.
Paralectotypes. INDIA: 1 3; 1 3, ‘Himalaya O.’ (MNHN, Paris); 1 2, Sikkim, Chasseurs Indigénes,
1894 (R. P. Bretandeau) (MNHN, Paris).
productus (Uroxys), 1933a : 389. Holotype 3, SouTH AMERICA: ‘Amer. aequinoct.’.
puncticollis (Panelus), 1931b : 406. Holotype [sex unknown], Sri LANKA: Kandy, vi.1908 (G. E. Bryant),
Paratype. Ski LANKA: 1 ex., same data as holotype.
quadridens (Pedaridium), 1932 : 225. Holotype [sex unknown], ARGENTINA: Entre Rios, Santa Elena,
30.1.1912 (G. E. Bryant).
rana (Onthophagus), 19316 : 268. Holotype 3, INDIA: Kodai Kanal (7. V. Campbell).
ranunculus (Onthophagus), 1913a : 461. LECTOTYPE 4, BRAZIL: Natal (W. H. Mann); here designated.
Paralectotypes. BRAZIL: 1 ¢, same data as lectotype; 5 2, Ceara, Baturite Mts (W. H. Mann); 1 3,
Para.
refulgens (Onthophagus), 1931b : 281. LECTOTYPE <3, Sri LANKA: Nuwara Eliya, 31.vii.1924; here
designated.
Paralectotypes. Sk1 LANKA: 1 3, 1 9, same data as lectotype; 1 9, Nuwara Eliya, 6234-8000 ft,
8-11.ii.1882 (G. Lewis); 2 3, 1 9, Ohiya, iv.1929. INDIA: 1 g, Kodai Kanal (7. V. Campbell); 1 3,
Madura, Shembaganur; 1 9, as previous but 1904-5 (P. de Breuil).
[Specimens from Sr1 LANKA: Nuwara Eliya, viii and xii (G. M. Henry) missing.]
regalis (Onthophagus), 1907c : 431. LECTOTYPE 4, Sri LANKA: here designated.
Paralectotype. Ski LANKA: 1 9, Yerbury.
[Specimen(s) from Sri LANKA: (E. E. Green) missing.]
rhinocerus (Liatongus), 1931b : 363. LECTOTYPE 4, INDIA: Kurseong, 4700-5000 ft, 25.vi.1910 (Annan-
dale); here designated.
Paralectotype. INDIA: 1 ¢ [not 9], Himalaya, Simla.
roberti (Pinotus), 1904 : 250 [original series: 2 3, 8 9]. LECTOTYPE 3, Brazi_: Chapada, 2600 ft,
xi.1902 (A. Roberts); here designated.
Paralectotypes. BRAZIL: 5 9, same data as lectotype ; 1 3, Para.
rotundus (Pycnopanelus), 1931b : 412. Holotype ? 3, INDIA: Madras.
106
rubripennis (Onthophagus), 1907c : 419. LECTOTYPE 4, Inp1A: Sikkim, environs de Kurseong (R. P.
de Bretandeau); here designated.
Paralectotypes. INDIA: 1 ¢, 1 9 (BMNH, London), 2 2 (MNHN, Paris), same data as lectotype ;
1 [sex unknown], Kurseong (R. P. de Bretandeau) (MNHN, Paris); 1 3, Sikkim.
rubripennis Arrow var. subcribratus (Onthophagus), see subcribratus.
ruficauda (Onthophagus), 1931b : 295. LECTOTYPE 6d, BurMa: Paungde (G. Q. Corbett); here desig-
nated.
Paralectotypes. BURMA: 1 3, same data as lectotype; 1 3, Toungoo (G. Q. Corbett); 1 2, Tharra-
waddy (G. Q. Corbett); 3 2, Mandalay (H. L. Andrewes).
runicus (Drepanocerus), 1909b: 93. Holotype 9, BURMA: Karen Mts, Asciuii Ghecu, 1400-1500 m,
iii-iv.1888 (L. Fea).
Paratypes. BURMA: 1 ¢ (BMNH), 2 3, 1 2 (MCSN, Genoa), same data as holotype.
setosus (Panelus), 1931b : 406. Holotype [sex unknown], Sri LANKA: Kandy, vi.1905 (G. E. Bryant).
sindensis (Gymnopleurus), 1931b : 66. Holotype 2, PAKISTAN: Karachi.
sladeni (Canthidium), 1904 : 249 [original series: 5]. LECTOTYPE 3, BraziL: Chapada, 2600 ft, xi.1902
(A. Robert); here designated.
Paralectotypes. BRAZIL: 1 g, 2 9, same data as lectotype.
socialis (Onthophagus), 1931b : 325. LECTOTYPE 4, Inpia: Nilgiri Hills (H. L. Andrewes); here desig-
nated.
Paralectotypes. INDIA: 12 3, 10 2, same data as lectotype ; 1 J, Bombay, Belgaum Div., Tavargatti,
ex dung, 31.x.1929 (B. M. Bhatia).
solidus (Onthophagus), 1931b : 265. Holotype 3, INDIA: Kumaon, Haldwani Dist. (H. G. Champion).
Current valid name: Onthophagus compactus Arrow (19336 : 421).
sternalis (Onthophagus), 1931b : 222. LECTOTYPE 4, INpIA: Berhampore; here designated.
Paralectotypes. INDIA: 1 3, 1 92, same data as lectotype; 1 g, 1 9, Fyzabad (R. W. G. Hingston).
[Specimens from InpIA: Murshidabad (E. T. Atkinson), Bihar, Pusa, vii (G. Bainbridge-Fletcher);
Chapra (Mackenzie); Dehra Dun, x (G. D. Pant); Bombay, Surat, vii (H. Maxwell Lefroy) missing.]
stevensi (Haroldius), 1931b : 416. Holotype [sex unknown], INDIA: Gopaldhara, Darjeeling, 4720-6100 ft,
4.ix.1914 (H. Stevens).
stevensi (Onthophagus laevis Harold var.), 19315: 172. LECTOTYPE 4, Inp1A: Sikkim, Gopaldhara,
Rungbong Valley (H. Stevens); here designated.
Paralectotypes. INDIA: 4 3, 5 2, same data as lectotype.
striatus (Delopleurus), 1931b: 411. LECTOTYPE [sex unknown], INDIA: United Prov. Dehra Dun,
2000 ft, vii.1927 (H. G. Champion); here designated.
[This specimen has been designated lectotype because it is uncertain that it is unique.]
subcribratus (Onthophagus rubripennis Arrow var.), 1907c : 420. LECTOTYPE 3, BurMaA: N. Chin Hills;
here designated.
Paralectotypes. INDIA: 1 9, Khasia Hills; 2 3, no locality label (E. J. C[antor]). BHUTAN: 1 @.
suillus (Onthophagus), 19316: 194. LECTOTYPE 3, INpDIA: South Mysore (H. L. Andrewes); here
designated.
Paralectotypes. INDIA: 1 3g, 2 9, same data as lectotype.
surdus (Copris), 19316: 132. LECTOTYPE 3, INpIA: Haldwani, Chakrata, ex dung, 3.v.1930 (B. M.
Bhatia); here designated.
Paralectotype. INDIA: 1 2, Kumaon, Tanakpur (H. G. C[hampion)).
symbioticus (Macropocopris), 1920: 437. Lectotype 3, AUSTRALIA: N. Queensland, off Wallaby
near vent, vii.1919 (Dr T. F. Illingworth); designated [as holotype] by Matthews (1972, Aust. J. Zool.
Suppl. 24 : 257).
Paralectotypes. AUSTRALIA: 13 3, 17 9, same data as lectotype.
Current valid name: Onthophagus symbioticus (Arrow) (Matthews, loc. cit.).
taprobanus (Onthophagus), 1931b: 299. LECTOTYPE 3, Sri LANKA: Bogawantalawa, 4900-5200 ft,
28.ii-12.iii.1882 (G. Lewis); here designated.
Paralectotypes. Ski LANKA: 1 92, same data as lectotype; 1 2 Dikoya, 3800-4200 ft, 6.xii.1881-
16.1.1882 (G. Lewis); 1 2 (G. Lewis).
107
tarantula (Sisyphus), 1909b : 92 [original series: 2 $]. LECTOTYPE 3, Sri LANKA; here designated.
Paralectotype. SRI LANKA: | 3, Bogawantalawa, 4900-5200 ft, 21.iii-4.iv.1882 (G. Lewis).
tarsius (Onthophagus), 1941 : 54. Holotype 3, NEw GuINeA: Japen I., R. Manai-Undei, 500 ft, x.1938
(L. E. Cheesman).
tibetanus (Onthophagus), 1907c : 428. LECTOTYPE 4, Cuina: Tibet, Brahmapoutra Valley, Chaksam,
12 000 ft, vii.1904 (H. J. Walton); here designated.
Paralectotypes. CHINA: 2 3, 6 9, same data as lectotype; 4 3, 3 9, Gyangtse, 13 000 ft, vi.1904
(H. J. Walton).
torticornis (Caccobius), 1931b : 144. LECTOTYPE 4, INDIA: Dehra Dun, cut grass, 10.vii.1922 (C. N.
Chatterjee); here designated.
Paralectotype. INDIA: 1 9, Dehra Dun, Nun Nadi, ex carrion, 17.vii.1922 (Dr M. Cameron).
[Specimen(s) from InpDIA: Almora Prov., Kali Valley, 5000 ft, vii (R. N. Parker) missing.]
triceratops (Onthophagus), 1913b : 195 [original series: 2 ¢]. LECTOTYPE 4, INp1A; here designated.
[1 3 specimen from INDIA: NE. Assam, Sadiya, 23.xi.1911 (Kemp) missing.]
tridenticeps (Chaeridium), 1913a:459. Holotype 3, BrAziL: Madeira-Mamore Railroad camp 28
(Mann & Baker).
trinitatis (Uroxys), 1933a : 391. LECTOTYPE 43, Trinipap: iii.1902 (G. E. Bryant); here designated.
Paralectotypes. TRINIDAD: 2 3, 5 2, same data as lectotype; 2 3, 2 9, 1903 (G. E. Bryant).
triste (Phacosoma), 19316 : 355 [original series: 2]. Holotype [sex unknown], INDIA: Nilgiri Hills (H. L.
Andrewes).
Paratype. INDIA: 1 ex., same data as holotype.
trochilus (Onthophagus), 1931b : 266. Holotype 3, BURMA: Tenasserim, Victoria Point (Doherty).
vincentiae (Uroxys), 1903 : 510 [original series: 2]. LECTOTYPE 4, St VINCENT: Cumberland (H. H.
Smith); here designated.
Paralectotype. St VINCENT: 1 92, same data as lectotype.
vividus (Onthophagus), 1907c : 428. LECTOTYPE 4, INpiA: S. India, only found in carcases; here
designated.
Paralectotypes. INDIA: 2 9, S. India; 7 3,7 2, Nilgiri Hills; 3 3,1 9, Nilgiri Hills (H. L. Andrewes);
1 3,1 2, Nilgiri Hills (G. F. Hampson); 2 3,2 92, Bangalore; 2 3, Trivandrum; 2 9, Malabar;.1 9,
India.
vulpinus (Onthophagus), 1931b : 262. LECTOTYPE 3, BuRMA: Momeit (Doherty); here designated.
Paralectotypes. BURMA: 1 3g, 1 9, Patkai Mts (Doherty).
vultur (Onthophagus), 1931b: 197. LECTOTYPE 4, INDIA: Bombay, Igatpuri, 2000 ft, 20-30.vi.1904;
here designated.
Paralectotype. INDIA: 1 2, Belgaum.
yunnanus (Synapsis), 1933b : 428. LECTOTYPE 3, CuinA: Yunnan, Tengyueh, 1909-10 (J. C. Brown);
here designated.
Paralectotypes. CHINA: 3 3, 1 9, same data as lectotype; 2 ¢, 1 2, Haut Yunnan, Tali.
[Specimen(s) from CuiNA: Chien Chuan Chou, 7700 ft, v (J. W. Gregory) missing.]
zebra (Onthophagus), 1931b : 249. LECTOTYPE 4, INpIA: Agra, Mhow (Selous); here designated.
Paralectotypes. INDIA: 1 3, 1 9°, Belgaum, Bombay.
LUCANIDAE
acutangulus (Dendroblax), 1935 : 122 [original series: 5]. LECTOTYPE 3, New ZEALAND: S. Island,
Greymouth (Helms); here designated.
Paralectotypes. NEw ZEALAND: 3 9, same data as lectotype.
Current valid name: Dendroblax earlei White (Holloway, 1961, Bull. Dom. Mus. Wellington 20 : 13).
acutangulus (Figulus), 1935 : 120. Syntypes, VIETNAM: 6 ex., Hoa Binh (A. de Cooman).
aratus (Figulus), 1935: 119. Syntypes, INDIA: 1 ex., Nilgiri Hills (H. L. Andrewes); 2 ex., Hattikeri,
under bark, 3.ii.1909 (H. L. Andrewes).
auritus (Aegotypus), 1943 : 142 [original series: 3 ¢]. LECTOTYPE <4, Borneo: here designated.
Paralectotypes. BoRNEO: 1 3, Kinabalu, 1904 (Rolle); 1 3, Kinabalu.
108
auritus (Figulus), 1927c : 63 [original series: 4]. Syntypes, WESTERN SAMOA: 2 ex., Apia, vi.-ix.1916
(Dr H. Swale); 1 ex., as previous but vi.1917; 1 ex., as previous but no date.
Current valid name: Figulus samoanus Kriesche (Arrow, 1935 : 119).
caeca (Vinsonella), 1940 : 93 [original series: 12]. LECTOTYPE 3, Mauritius: Mt Cocotte, 26.xi.1938
(J. Vinsom), here designated.
Paralectotypes. MAURITIUS: 2 3 (BMNH, London), 1 2 (MNHN, Paris) same data as lectotype.
[It is possible that there is one more male paralectotype in MNHN, Paris, but it was not possible to
check at the time that this catalogue was compiled.]
carolinensis (Dorcus), 1939a : 84. LECTOTYPE 3, CAROLINE ISLANDS: Ponape I., Kolonie, 9.iii.1936
(Z. Ono), here designated.
Paralectotypes. CAROLINE ISLANDS: 1 3, same data as lectotype but 2.ii.1936 (BPBM, Honolulu);
1 2, Ponape I., 29.ii.1936 (Z. Ono) (BPBM, Honolulu); 1 3, Ponape I., 6.iii.1936 (Z. Ono); 1 3,1 9,
Ponape I., Tamataman-Sakir [not Yamataman], 20.iii.1936 (S. Otomo); 1 3, Palao I., 8.iv.1936 (Z.
Ono).
cheesmani (Figulus), 1938 : 61. Holotype [sex unknown], New Guinea: Papua, Kokoda, 1200 ft, viii.1933
(L. E. Cheesman).
concatenatus (Figulus), 1938 : 59. Holotype [sex unknown], BORNEO: W. Sarawak, Mt Matang, 1000 ft,
3.xi.1914 (G. E. Bryant).
coomani (Figulus), 1935: 120. Syntypes, VIETNAM: 2 ex., (BMNH, London), 1 ex., (MNHN, Paris),
Backau, 1907-8 (P. Lemée); 7 ex., (BMNH, London), 1 ex., (MNHN, Paris), Hoa Binh (A. de Cooman).
delicatus (Aegus), 1943 : 141 [original series: 2 $]. LECTOTYPE 3, West MataysiA: Perak (Doherty);
here designated.
Paralectotype. WesT MALAysIA: 1 3, same data as lectotype.
dublonensis (Dorcus), 1939b : 580. Holotype 3, CAROLINE ISLANDS: Truk Islands, Dublon I., 23.xii.1935
(Z. Ono).
Paratype. CAROLINE ISLANDS: 1 2, Truk Islands, Dublon I., 5.i.1936 (Z. Ono).
elongatus (Penichrolucanus), 1935 : 122. Holotype 3, WesT MALAysIA: Selangor, Kuala Lumpur, Gombak
Valley, 13.x.1921 (H. M. Pendlebury).
frater (Aegus), 1935:114. Holotype 3, MoLuccas: Halmahera, Gamkonora, 1200 m, iv.1931 (G.
Heinrich).
furcifer (Lucanus) 1949 : 46. LECTOTYPE 3, CuinA: Yunnan; here designated.
Paralectotypes. CHINA: 1 3, 1 9, same data as lectotype; 1 g, 1 2, Szechuan, ouy Sy, 1911 (R. P.
Mombelg). INDIA: 1 3, Sikkim, Lachen Lochung, viii (MNHN, Paris).
[A lectotype has been designated because although Arrow says that the type is from Yunnan, no
specimen was so labelled.]
henryi (Dorcus), 1935 : 110. Holotype 3, Sri LANKA: Trincomali Dist., Pulmoddai, 10.viii.1934 (G. M-
Henry).
histrio (Dorcus), 1935: 109. LECTOTYPE 3, INDIA: Kumali Hill, 29.ii.1916 (K. Govindaraj); here
designated.
Paralectotype. INDIA: 1 3, Coorg, Periambadi Ghat, 28.v.1917 (7. R. Rao).
humilis (Dorcus), 1935 : 111 [original series: 7]. LECTOTYPE 3, Inp1A: Sikkim, Gopaldhara, Rungbong
Valley, 1911 (H. Stevens); here designated.
Paralectotypes. INDIA: 1 9, Sikkim, Gopaldhara, Rungbong Valley, Darjeeling, 1911 (W. K. Webb).
BHUTAN: 1 9, Pedong, 1899 (L. Durel); 1 ¢ (BMNH, London), 1 3, 1 2 (MNHN, Paris), same as
previous but 1913.
immundus (Dorcus), 1938 : 56 [original series: 3 3, 1 2]. Holotype 3, INDIA: Coimbatore Dist., Camp
Valparai, 3500 ft, 1937.
Paratypes. INDIA: 1 ¢ (BMNH, London), 1 ¢ (MNHN, Paris), same data as holotype.
[1 9, same data as holotype (MNHN, Paris) missing.]
leveri (Penichrolucanus), 1938 : 61 [original series: 3]. Syntypes, SOLOMON ISLANDS: 1 ex., Mamowa,
rotten wood, xi.1931 (R. A. Lever); 2 ex., as previous but Guadalcanal, Mamawa.
marginatus (Aegotypus), 1935 : 115. Holotype 2, BORNEO: ex. coll Parry.
nageli (Dorcus), 1935 : 112. Holotype 3, INDIA: Assam (P. Nage/).
109
nicobaricus (Penichrolucanus), 1935 : 123. Holotype [sex unknown], NicoBAR IsLANDs: (MNHN, Paris).
oberthuri (Torynognathus), 1935: 116. LECTOTYPE 3, SUMATRA: Semangoes Forest (J. Bouchard);
here designated.
Paralectotypes. SUMATRA: 1 3, 3 9, 1 [sex unknown, head missing], (BMNH, London), 24 [sex
unknown] (MNHN, Paris), 2 [sex unknown] (Coll. M. Bomans), same data as lectotype.
reticulatus (Cardanus), 1943 : 140. Holotype [sex unknown], SUMATRA: Sungei Kumbang, Korinchi,
4500 ft, iv.1914.
rhodesianus (Nigidius), 1935 : 119. Syntypes, RHODESIA: 4 ex. (BMNH, London), 1 ex. (MNHN, Paris),
Salisbury (Marshall); 1 ex., Mashonaland, Salisbury, xi.1900 (G. A. K. Marshall); 1 ex., as previous
but iii.1901; 1 ex., Kashitu, N. of Broken Hill, i.1905 (H. C. Dollman); 1 ex., as previous but ii.1905.
Current valid name: Nigidius rhodesianus Peringuey (Arrow, 1938 : 53).
scorpio (Platyfigulus), 1935: 117. Holotype 3, locality unknown [‘The unique specimen, which has
generously been presented to the British Museum by the authorities of the Colombo Museum, is of
unknown origin, but must be assumed to be a native of Ceylon’, Arrow (loc. cit.)].
serratus (Figulus), 1938 : 60 [original series: 3]. Syntypes, BORNEO: 1 ex., Sarawak, Retuh, 14.v.1914
(G. E. Bryant); 1 ex., Sarawak, Tabehang, 12.v.1914 (G. E. Bryant).
sumatrensis (Penichrolucanus), 1935: 124. Holotype [sex unknown], SUMATRA: Palembang (MNHN,
Paris).
swalei (Aegus), 1927c: 61 [original series: 2]. LECTOTYPE 3, WESTERN SAMOA: Apia, vi.-ix.1916
(Dr H. Swale); here designated.
Paralectotype. SAMOA: 1 2 (Dr H. Swale).
[The ° paralectotype is 21 mm long, not 11 mm.]
tutuilensis (Aegus), 1927c : 62 [original series: 3]. LECTOTYPE 3, AMERICAN SAMOA: Tutuila, Leone
Road, 9.vii.1923 (Sweezey & Wilder); here designated (BPBM, Honolulu).
Paralectotypes. AMERICAN SAMOA: 1 3 (BMNH, London), 1 ¢ (BPBM, Honolulu), Tutuila, 18.xii.
(H. C. Kellers).
upoluensis (Aegus), 1927c : 60. LECTOTYPE <3, ‘SAMoa’: Navigator Is.; here designated.
Paralectotypes. ‘SAMOA’: 1 ¢ (S. J. Whitmee) [collector from BMNH Coleoptera accessions register].
WALLIs ISLANDS: 1 9. WESTERN SAMOA: 1 3, Lotopa, 29.iv.1917 (H. Swale); 1 3, as previous but
1.111.1917; 1 3, Apia, vi-ix.1916 (H. Swale); 1 3, ‘Samoa’, 20.v.1917 (H. Swale); 1 3, as previous but
no date.
[Aegus upoluensis was proposed by Arrow as a ‘nom.n.’ for Alcinus dilatatus Fairmaire sensu
Waterhouse (Trans. ent. Soc. Lond. 1875 : 163). Waterhouse’s description was based on a misidentifica-
tion and Arrow’s note is, therefore, a valid species description.
The lectotype and the specimen collected by Whitmee are the two specimens described by Waterhouse
and, although they are not actually mentioned in Arrow’s description, it is inferred that he has seen
them. Also, the only identification label on the whole syntype series is Arrow’s label ‘Aegus upoluensis
Arrow, Type’ on the major male described by Waterhouse. This specimen is therefore designated
lectotype.]
ursulus (Dorcus), 1938 : 55. Holotype 3, BHUTAN: Pedong, 1913.x. (L. Durel).
Paratypes. [1 3, 1 2 in Oberthiir Collection (MNHN, Paris) missing.]
variolosus (Cardanus), 1935 : 121 [original series: 2]. Holotype [sex unknown], THAILAND: (Bates).
Paratype. INDIA: 1 ex., Assam, Rotung, 1400 ft, ex rotten wood, 23.xii.1911 (S. W. Kemp) (ZSI,
Calcutta).
vernicatus (Dorcus), 1938: 58. Holotype 3, INDIA: Assam, Shillong District, Oberthiir Collection
(MNHN, Paris) [missing].
wardi (Dorcus), 1943: 136. LECTOTYPE <3, Cuina: SE. Tibet, Zayul 1000-1200 ft, vi & vii.1935
(R. J. H. Kaulback); here designated.
Paralectotypes. CHINA: 1 9, SE. Tibet, Zayul, Di Chu Valley, 11 000 ft, 23.viii.1933 (F. Kingdon-
Ward & R. J. H. Kaulback). BuRMA: 1 92, Seinghku Valley, 28°9’ N, 97°25’ E, 6.vi.1926 (F. Kingdon-
Ward).
PASSALIDAE
affinis (Eumelosomus), 1907a : 465. Syntypes. UGANDA: 4 ex., E. Ruwenzori, 600-1200 ft, 1906 (Hon.
G. R. Legge & A. F. R. Wollaston).
110
antillarum (Neleides), 1907a : 452. Syntypes. GRENADA: 3 ex., Chantilly Estate (windward side) (H. H.
Smith); 2 ex., Grand Etang (windward side), 1900 ft (H. H. Smith); 3 ex., Windsor (H. H. Smith);
1 ex., Windsor Estate (windward), 500 ft, 28.iii, wooded ravine, under a rotten log. Found at same
place and are probably of the same species [then in different handwriting], i.e. as larvae.
archidonae (Petrejus), 1907a : 465. Holotype [sex unknown], Ecuapor: Archidonae.
borealis (Chilomazus), 1907a : 467. Holotype [sex unknown], INDIA: Assam, Nagas (Doherty).
centralis (Tetraracus), 1907a : 458 [original series: 2]. Syntypes, Costa RICA: 2 ex., Volcan de Barba
1600 m (P. Biolley).
congoensis (Didymus), 1907a : 463. Syntypes, ANGOLA: 4 ex., San Salvador. ZAIRE: 2 ex., R. Congo,
Mayanda.
crassus (Didymus), 1907a : 465. Holotype, UGANDA: E. Ruwenzori, 6000-13 000 ft, 1906 (Hon. G. Legge
& A. F. R. Wollaston).
curvilineatus (Didymus), 1907a : 462. Syntypes, UGANDA: 1 ex., Msozi, 4300 ft (Delme Ratcliffe); 2 ex.,
Msozi, 11.1903 (Delme Radcliffe). UGANDA: 1 ex., Kavirondo.
isthmicus (Veturius), 1907a : 453. Syntypes, Costa RICA: 1 ex., PANAMA: 1 ex., Bugaba (Champion).
laevisternus (Didymus), 1907a : 463. Syntypes, KENYA: 5 ex., Kikuyu.
nasutus (Eumelus), 1907a : 459. Syntypes, EcuApor: 1 ex., Cachabe, low c[amp ?], xi.1896 (Rosenberg);
2 ex., 1882-3 (Siemradski); 1 ex., as previous, no date.
[Specimens collected by Siemradski (MNHN, Paris) missing.]
peruvianus (Petrejus), 1907a : 456. Syntypes, PERU: 4 ex.
peruvianus (Veturius), 1907a : 455. Syntypes, PERU, 2 ex., Marcapata R. (J. Kalinowski); 1 ex., Peru.
platypleura (Erionomus), 1907a : 461. Syntypes, KENYA: 6 ex., Kavirondo.
pulchellus (Leptaulacides), 1907a : 466. Holotype [sex unknown], New Guinea: Ekeikei, iv.1900 (Pratt).
punctatostriatus (Veturius), 1907a : 454. Syntypes, GUYANA: 5 ex., Demarara, Georgetown, 1879 (W.
Harper).
ruwenzoricus (Didymus), 1907a : 464. Syntypes, UGANDA: 6 ex., E. Ruwenzori, 6000-13 000 ft, 1906
(Hon. G. Legge & A. F. R. Wollaston).
solidus (Triaenurgus), 1907a: 452 [original series: 2]. Syntypes, GUATEMALA: 2 ex., Quezaetenango
Chuipache (Richardson).
spinosus (Petrejus), 1907a : 457 [original series: 4]. Syntypes, ECUADOR: 4 ex., Cachabe, Low C[amp 7],
1896 (Rosenberg).
TROGIDAE
acutipes (Cloeotus), 1903 : 519. Syntypes, BRAZIL: 9 ex., Rio de Janeiro (Fry); 3 ex., Brazil. Costa RICA:
2 ex., Irazu, 600-700 ft (H. Rogers).
birmanicus (Trox), 1927b : 467. LECTOTYPE 3, BurMA: Prome; here designated.
Paralectotypes. BURMA: 1 ex., Pakokku, 180 ft, ix.1915 (Miss Molesworth); 1 ex., Burma; 1 ex.,
Minhla, 1881 (Comotto) (MCSN, Genoa).
[Specimen(s) from CaAmMBoDIA: Kompong Kedey, v (R. V. de Salvaza) missing.]
[The statement ‘type (3) in coll. British Museum’ by Haaf (1954, Ent. Arb. Mus. Georg Frey 5 : 389)
is incorrect. Although labelled as ‘type’ by Arrow, the specimen was not designated as the holotype by
him. It is this specimen which has been selected as the lectotype.]
bugabensis (Cloeotus), 1903 : 519 [original series: 2]. Syntypes, PANAMA: 2 ex., Bugaba, 800-1500 ft
(Champion). [One of these syntypes is also a syntype of Cloeotus viridipennis Bates (Arrow, loc. cit.).]
crassicollis (Cloeotus), 1093 : 518 [original series: 2]. Syntypes, ST VINCENT: 1 ex., Forest Morne a Garon,
1500 ft, in rotten wood, 27.x; 1 ex., (H. H. Smith).
expansus (Trox), 1900: 22. Holotype, SoMALI REPUBLIC: central & east Somaliland, 5.vi.-29.x.1897
(C. A. V. Peel).
Sryi (Cryptogenius), 1909c : 501. Holotype, BRAZIL: Rio de Janeiro (Fry).
lobicollis (Trox), 1927b : 466. Syntypes, BURMA: 5 ex. (BMNH, London) 3 ex. (IP, Eberswalde), 2 ex.
(MCSN, Genoa) Tenasserim, Moulmein, v.1887 (Fea); 1 ex., Burma.
[Specimens from BuRMA: Rangoon, vi and x (Fea) missing.]
111
mollis (Trox),1927b : 468. Syntypes, BORNEO: | ex., Borneo; 1 ex., Medan (L. Fulmek) (IP, Eberswalde);
SUMATRA: | ex., Palembang, 1900 (J. Schmitz).
[Specimen(s) from West MALAysIA: Selangor, Kuala Lumpur, viii. (H. M. Pendlebury) missing.]
rufopiceus (Cloeotus), 1903 : 517. Syntypes, St VINCENT: 2 ex., (H. H. Smith); GRENADA: 1 ex., Grand
Etang (windward side), 1900 ft (H. H. Smith).
testudo (Trox), 1927b : 466. LECTOTYPE <3, PAKISTAN: Karachi; here designated.
Paralectotypes. PAKISTAN: 2 ex., same data as lectotype; 1 ex., ‘Ind. bor.’.
[The statement ‘3-Typus in coll. British Museum, London’ by Haaf, (1954, Ent. Arb. Mus. Georg
Frey 5 : 365) is incorrect. Although labelled as ‘Type’ by Arrow, the specimen was not designated as
the holotype by him. It is this specimen which has been selected as the lectotype.]
HYBOSORIDAE
acuticollis (Phaeochroops), 1907c : 436. Holotype, BoRNEO: Kinabalu.
amplus (Phaeochrous), 1909c : 499. LECTOTYPE 3, CAMEROUN: Mundame (R. Rohde); here designated.
Paralectotypes. CAMEROUN: 1 3g, 1 2 (BMNH, London), 6 3g, 1 9 (IP, Eberswalde), same data as
lectotype.
arabicus (Phaeochrous), 1909c : 497. LECTOTYPE 3, YEMEN: (Millingen); here designated.
Paralectotypes. YEMEN: 6 3, 1 92, same data as lectotype.
batuensis (Phaeochroops), 1909c : 495. Syntypes, SULAWESI: 6 ex. (BMNH, London), 1 ex. (MCSN,
Genoa), Batu I., 1896-7 (H. Raap).
camerunensis (Phaeochrous), 1909c : 499. LECTOTYPE 3, CAMEROUN: Mundame (R. Rohde); here
designated.
Paralectotypes. CAMEROUN: 1 3g, 1 2 (BMNH, London), 5 3, 4 2 (IP, Eberswalde), same data as
lectotype.
cinereicollis (Phaeochridius), 1942 : 924. LECTOTYPE 4, Borneo: Sarawak, foot of Mt Dulit, junction
of rivers Tinjar & Lejok, 2.x.1932, trap 2, goat (B. M. Hobby & A. W. Moore); here designated.
Paralectotypes. BORNEO: 2 3, same data as lectotype; 1 3, same data as lectotype but 1.x.1932;
1 3, same data as lectotype but 20.ix.1932, trap 1, fish; 2 3, same data as lectotype but 22.ix.1932,
trap 1, fish; 1 3, same data as lectotype but 25.ix.1932, trap 1, fish; 1 3, same data as lectotype but
4.x.1932, trap 1, fish; 1 g, same data as lectotype but 4.ix.1932, old secondary forest; 1 3, same data as
lectotype but 24.ix.1932, light-traps 6 & 7.
crassicollis (Liparochrus), 1925 : 329. LECTOTYPE [sex unknown], New Guinea: Kani Mts (Bennigsen);
here designated.
[Although the sex of the lectotype cannot be determined without dissection, it has been so designated
because the syntypes from NEw GuINEA: Mamberano River, Pioneerbivak, vii—viii and xii-i (W. C.
van Heurn) cannot be found in the collection of IP, Eberswalde.
dissimilis (Phaeochrous), 1909c : 496. LECTOTYPE 3, BurMA: Tenasserim, Moulmein, v.1887 (Fea);
here designated.
Paralectotypes. BURMA: 1 3, 4 2 (BMNH, London), 4 3, 2 2 (MCSN, Genoa), 1 3, 1 2 (MHNH,
Paris), 4 3, 2 2 (SMT, Dresden), same data as lectotype.
dux (Liparochrus), 1909c : 489. LECTOTYPE 3, New Guinea: Moroka, 1300 m, vii—xi.1892 (Loria);
here designated.
Paralectotypes. NEw GUINEA: 1 3, same data as lectotype (MCSN, Genoa); 1 9, Paumomu R.
ix—xii.1892 (Loria).
exaratus (Chaetodus), 1909c : 491. Syntypes, BRAZIL: 1 ex., Rio de Janeiro [Fry collection but doubtfully
collected by Fry]; 1 ex., ‘Adelops striatus mihi, L in Brasilia (Dr. Sommer)’.
fumipennis (Coelodes), 1909c : 491. LECTOTYPE 4, BraziL: Ega; here designated.
Paralectotypes. BRAZIL: 1 9, same data as lectotype; 2 9, Amazon (Bates); 1 3,2 9, Para.
gigas (Phaeochroops), 1907c : 436. Syntypes, BORNEO: 2 ex., Pengaron (Doherty).
[Specimens from West Ma aysIA: Perak (Doherty) missing.]
indicus (Phaeochroops), 1907c : 359 [original series: 3]. Syntypes, INDIA: 3 ex., Nigiri Hills (H. L.
Andrewes).
laetus (Microphaeochroops), 1942 : 926. Holotype, BORNEO: Sarawak, Mt Dulit, 4000 ft, primitive moss
forest, 26.x.1932 (B. M. Hobby & A. W. Moore).
112
lugubris (Liparochrus), 1925: 328. LECTOTYPE 3, New Guinea: Mamberamo R., Pioneerbivak,
xii. 1920 (W. C. van Heurn); here designated.
Paralectotype. NEw GuINEA: | 3, same data as lectotype (IP, Eberswalde).
[The female specimen cannot be traced.]
mashunus (Phaeochrous), 1909c : 498. LECTOTYPE 3, RuHopesiA: Mashonaland, Salisbury, xi.1900
(G. A. K. Marshall), here designated.
Paralectotypes. RHODESIA: | 3, same data as lectotype but xii. 1904: 15 3g, 10 2 (BMNH, London),
3 g, 2 2 (MCSN, Genoa), 3 (sex unknown) (MNHU, Berlin), 4 3, 1 2 (SMT, Dresden), same data as
lectotype but no date; 1 2, Mashonaland. MOZAMBIQUE: 4 3, 1 2 Nyassa (Thelwall).
mentaweiensis (Phaeochroops), 1909c : 495. Syntypes, MENTAWEI ISLANDs (Indonesia): 2 ex. (BMNH,
London), 3 ex. (MCSN, Genoa), Sipora, Sereinu, v—vi.1894 (Modigliani).
Current valid name: Phaeochroops rattus Arrow (Arrow, 1942 : 927).
nanus (Phaeochrous), 1942 : 924 [original series: ‘over 50°]. LECTOTYPE 3, Inp1A: Bihar, Pusa, 24.vi.
1924 (Mukerjee); here designated.
Paralectotypes. INDIA: 2 3, same data as lectotype; 1 9, same data as lectotype but 27.vi.1916
(Fletcher); 2 2, same data as lectotype but 1.viii.1916 (D. Nandan); 1 3, as previous but 3.viii.1916;
1 9, Bihar, Pusa, 18.vi.1916, at light (H[rest illegible]); 1 2, as previous but no date; 1 3, 6 9, Bengal,
Chapra (Mackenzie).
niasianus (Phaeochroops), 1909c : 494. Syntypes, Nias I. (Indonesia): 3 ex. (BMNH, London), 5 ex.
(MCSN, Genoa), 1897-8 (U. [not H.] Raap).
nigripennis (Coelodes), 1903: 516. LECTOTYPE 3d, St VINCENT: leeward side (H. H. Smith); here
designated.
nitidus (Phaeochrous), 1909c : 497. LECTOTYPE 3, TANZANIA: Masai; here designated.
Paralectotypes. TANZANIA: 2 3g, 2 9, same data as lectotype; 2 3, 2 92, Kilimanjaro; 1 9, ‘Germ.
E. Afr.’ (Rev. A. N. Wood).
obscurus (Hybochaetodus), 1909c : 500 [original series: 2]. LECTOTYPE [sex unknown], PERu: Vilcanota;
here designated.
[Although the sex of the lectotype cannot be determined without dissection, it has been so designated
because the other syntype cannot be found in the collection of MNHU, Berlin.]
opacicollis (Phaeochroops), 1909c : 493. Syntypes, BURMA: 1 ex. (BMNH, London), 1 ex. (MCSN,
Genoa), Tenasserim, Plapoo, iv.1887 (Fea); 1 ex., Tenasserim, Mt Moolevit, 1000-1 ?00 m, iv.1887
(Fea) (MCSN, Genoa).
pallidus (Phaeochrous), 1909c : 496. LECTOTYPE 4, INDIA: South Mysore (H. L. Andrewes); here
designated.
Paralectotypes. INDIA: 5 3, same data as lectotype; 3 3, Nilgiri Hills (G. F. Hampson). Sr1 LANKA:
Pigs dove
peninsularis (Microphaeochroops), 1942: 925 [original series: 3]. Syntypes, WeEsT MALAYSIA: 3 ex.,
Perak, Jor Camp, 2000 ft, 25.viii.1922 (E. Seimund).
peninsularis (Phaeochroops), 1909c : 492 [original series: 1 3, 1 9]. LECTOTYPE 3, West MALaysIA:
Perak (Doherty); here designated.
Paralectotype. WEsT MALAysIA: 1 9, same data as lectotype.
punctatum (Dicraedon), 1911 : 396 [original series: 3 9]. LECTOTYPE 9, GUATEMALA: Zapote
(Champion), here designated.
Paralectotypes. COLOMBIA: 1 9, Cali, ix—xii.1894 (W. Rosenberg); 1 9°, same data as previous but
9-10.1.1894 (SMT, Dresden).
punctipennis (Coelodes), 1909c : 491. LECTOTYPE 3, Ecuapor: Mirador; here designated.
Paralectotypes. Ecuapor: 1 3, same data as lectotype; 1 2, Canelos (Buckley); 1 9, Ecuador. PERU:
1 9, Amazon, Nauta.
punctulatus (Phaeochroops), 1942: 926. Syntypes, BORNEO: 1 ex., Sarawak, Matang, 3-13.vii.1909
(C. J. Brooks); 1 ex., Sarawak, Mt Merinjak, 2000 ft, 26.v.1914 (G. E. Bryant); 1 ex., Sarawak, foot of
Mt Dulit, junction of rivers Tinjar & Lejok, trap 1, fish, 25.ix.1932 (B. M. Hobby & A. W. Moore);
1 ex., Sarawak, Mt Kulalong, Long Manian, 8.xi.1932 (B. M. Hobby & A. W. Moore); 3 ex., Kinabalu.
pusillus (Hapalonychus), 1911 : 396 [original series: 2]. Holotype 3, PARAGUAY.
Paratype. PARAGUAY: 1 ¢ (Dr Bohls) (SMT, Dresden).
113
rattus (Phaeochroops), 1909c : 494. Syntypes, SUMATRA: 2 ex. (BMNH, London), 4 ex. (MCSN, Genoa),
Setinjak, 1800 ft, 1.1898; 2 ex., Si-Rambé, xii.1890-iii.1991 (E. Modigliani).
thomensis (Phaeochrous), 1909c : 499. LECTOTYPE 3, SAo Tomé: Vista Alegre, 200-300 m, x.1900
(L. Fea); here designated.
Paralectotypes. SAo TomE: 1 3, 1 9, same data as lectotype (MCSN, Genoa).
timidus (Liparochrus), 1909c : 490. LECTOTYPE 6d, AustrALiA: Alexandria (W. Stalker); here desig-
nated.
Paralectotypes. AUSTRALIA: 8 ex. (BMNH, London), 1 ex. (ANIC, Canberra), 1 ex. (MCSN, Genoa),
same data as lectotype; 6 ex. (BMNH, London), 1 ex. (MCSN, Genoa), same data as lectotype but
x.1905; 8 ex. (BMNH, London), 1 ex. (MNHU, Berlin), 2 ex. (MCSN, Genoa) same data as lectotype
but 30.xii.1905; 1 ex., same data as lectotype but 25.ii.1906 (MCSN, Genoa); 14 ex. (BMNH, London),
1 ex. (ANIC, Canberra), same data as lectotype but 13-20.iii.1905; 2 ex., same data as lectotype but
20.11.1906 (MNHU, Berlin).
Current valid name: Liparochrus dilatifrons Blackburn (Arrow, 1925 : 328).
uniformis (Phaeochridius), 1925 : 330. LECTOTYPE 3, Borneo: Kinabalu; here designated.
Paralectotype. BORNEO: 1 9, same data as lectotype (IP, Eberswalde).
vulpecula (Phaeochroops), 1909c : 493. Syntypes, MENTAWEI ISLANDs (Indonesia) : 2 ex. (BMNH, London),
3 ex. (MCSN, Genoa) Sipora, Sereinu, v—vi.1894 (Modigliani).
GEOTRUPIDAE
bahiae (Athyreus),1913a: 463, Syntypes, BRAZIL: 2 ex., Bahia; 1 ex., Goyas Prov, Jatahy; 1 ex., Itaparica
Is, 28.xii.1906 (E. G. B. Meade-Waldo).
fissicollis (Athyreus), 1913a : 462. Holotype [sex unknown], BRAZIL: Natal (W. H. Mann).
loweri Blackburn var. /aratinum (Bolboceras), see laratinum.
laratinum (Bolboceras loweri Blackburn var.), 1916 : 493 [original series: 2]. Syntype [sex unknown],
LaraT I. (Indonesia) (F. Muir).
parcepunctatum (Bolboceras), 1913a : 461. Holotype 3, BRAZIL; Bahia.
Paratypes. BRAZIL: 1 3, 4 9, same data as holotype; 1 9, Ceara-Mirim, Rio Grande de Norte
(W. M. Mann); 1 9, Indepencia, Parahyba (Mann & Heath).
References
Arrow, G,. J. 1900. Jn Gahan, C. J. & Arrow, G. J., On a collection of insects and arachnids made in
1895 and 1897 by Mr C. V. A. Peel, F.Z.S., in Somaliland, with descriptions of new species. 5. Cole-
optera. Proc. zool. Soc. Lond. 1900 : 21-33, pl. 1.
— 1903. On the laparostict Lamellicorn Coleoptera of Grenada and St. Vincent (West Indies). Trans.
ent. Soc. Lond. 1903 : 509-520, 4 figs.
— 1904. In Gahan, C. J. & Arrow, G. J., List of the Coleoptera collected by Mr. A. Robert at Chapada,
Matto Grosso (Percy Sladen expedition to central Brazil). Proc. zool. Soc. Lond. 1903(2) : 244-258,
pl. 28.
1906. On Lamellicorn Coleoptera from Portuguese West Africa, with descriptions of new species.
Ann. Mag. nat. Hist. (7) 18 : 127-136, 1 fig.
1907a. A contribution to the classification of the coleopterous family Passalidae. Trans. ent. Soc.
Lond. 1906 : 441-469.
—— 1907b. Some new species and genera of Lamellicorn Coleoptera from the Indian empire (part 1).
Ann. Mag. nat. Hist. (7) 19 : 347-359.
—— 1907c. Some new species and genera of Lamellicorn Coleoptera from the Indian empire (part 2).
Ann. Mag. nat. Hist. (7) 19 : 416-439, 3 figs.
— 1908. Notes on the coleopterous genus Onitice/lus and descriptions of some new species from India.
Ann. Mag. nat. Hist. (8) 1: 178-183.
—— 1909a. On some new species of Coleoptera from Rhodesia and adjacent territories. Ann. Mag. nat.
Hist. (8) 3 : 517-523.
—— 1909b. Four new Lamellicorn Coleoptera from the Oriental Region. Ann. Mag. nat. Hist. (8) 4 : 91-94.
—— 1909c. On the characters and relationships of the less-known groups of Lamellicorn Coleoptera,
with descriptions of new species of Hybosorinae etc. Trans. ent. Soc. Lond. 1909 : 479-507.
114
— 1911. On Lamellicorn beetles belonging to the subfamilies Ochodaeinae, Orphninae, Hybosorinae
and Troginae. Ann. Mag. nat. Hist. (8) 7 : 390-397.
—— 1913a. Some new species of Lamellicorn beetles from Brazil. Ann. Mag. nat. Hist. (8) 11 : 456-466.
— 1913b. Zoological results of the Abor expedition 1911-12, Coleoptera IV. Lamellicornia. Rec.
Indian Mus. 8 : 191-196.
—— 1916. On the Lamellicorn Coleoptera of Larat Island. Ann. Mag. nat. Hist. (8) 18 : 492-498.
—— 1919. A remarkable new ball-rolling beetle (family Scarabaeidae). Ann. Mag. nat. Hist. (9) 3: 433-
435, 1 fig.
—— 1920. A peculiar new genus of Australian beetles. Ann. Mag. nat. Hist. (9) 6 : 434-437.
—— 1925. Notes on the coleopterous families Hybosoridae and Trogidae. Ann. Mag. nat. Hist. (9)
15 : 328-331.
—— 1927a. Notes on the coleopterous genus Sisyphus. Ann. Mag. nat. Hist. (9) 19 : 456-465, pl. 12.
—— 1927b. A note on the coleopterous genus 7rox, with descriptions of a few new Asiatic species. Ann.
Mag. nat. Hist. (9) 19 : 465-468.
——1927c. Coleoptera, Clavicornia and Lamellicornia. Insects Samoa. Pt. 4, fasc. 1 : 35-66, 13 figs.
— 193la. The coleopterous genus Trichillum (Copridae), with a key to the species. Ann. Mag. nat.
Hist. (10) 8 : 609-611.
— 1931b. Fauna Br. India. Coleoptera, Lamellicornia part 3 (Coprinae). xii+428 pp., 13 pls, 1 map,
60 figs.
—— 1932. New species of Lamellicorn beetles (subfamily Coprinae) from South America. Stylops 1 : 223-
226, 3 figs.
—— 1933a. The genus Uroxys (Coleoptera, Copridae), with descriptions of some new species. Ann.
Mag. nat. Hist. (10) 11 : 385-399, 9 figs.
—— 1933b. Notes on Coprid Coleoptera, with descriptions of a new genus and a few new species. Ann.
Mag. nat. Hist. (10) 12 : 421-430, 4 figs.
—— 1935. A contribution to the classification of the coleopterous family Lucanidae. Trans. R. ent
Soc. Lond. 83 : 105-125, pl. 4, 5 figs.
1938. Some notes on stag-beetles (Lucanidae) and descriptions of a few new species. Ann. Mag.
nat. Hist. (11) 2 : 49-63, pl. 4.
1939a. Three new species of Lamellicorn beetles from the Caroline Islands. Entomologist’s mon.
Mag. 75 : 84-87.
—— 1939b. The Lucanid Coleoptera of the Caroline Islands. Ann. Mag. nat. Hist. (11) 4: 579-582, pl. 4.
—— 1940. A new genus of blind beetles of the family Lucanidae (Col.). Proc. R. ent. Soc. Lond. (B)
9 : 93-96, 11 figs.
— 1941. A few new species of the genus Onthophagus (Coleoptera, Copridae) from New Guinea and
Japen Island. Ann. Mag. nat. Hist. (11) 8 : 48-55.
—— 1942. A few new species of the Scarabaeid subfamily Hybosorinae (Coleoptera), with a key to the
genus Phaeochroops. Ann. Mag. nat. Hist. (11) 9 : 923-928.
—— 1943. On the genera and nomenclature of the Lucanid Coleoptera, and descriptions of a few new
species. Proc. R. ent. Soc. Lond. (B) 12 : 133-143.
— 1949. Fauna Br. India. Coleoptera, Lamellicornia part 4 (Lucanidae and Passalidae). xi+ 274 pp.,
23 pls.
Bacchus, M. E. 1974. A catalogue of the type-specimens of the Cetoniinae (Coleoptera : Scarabaeidae)
described by G. J. Arrow, with a complete bibliography of his entomological works. Bull Br. Mus.
nat. Hist. (Ent.) 31 : 23-44.
115
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Bulletin of the
A revision of the Neotropical wasp genus
Trigonopsis Perty (Hymenoptera: Sphecidae)
C. R. Vardy
Entomology series Vol 37 No4 27 July 1978
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© Trustees of the British Museum (Natural History), 1978
ISSN 0524-6431 Entomology series
Vol 37 No 4 pp 117-152 —
British Museum (Natural History)
Cromwell Road |
London SW7 5BD ' Issued 27 July 1978.
A revision of the Neotropical wasp genus T. rigonopsis
Perty (Hymenoptera : Sphecidae)
C. R. Vardy.,.
Department of Entomology, British Museum (Natural History), Cromwell Road, robe
SW7 5BD
Contents
Synopsis. : ; ; F ; : d ; : : : ' Een hy
Introduction . . : : , ; : : ; . : : eee
Acknowledgements . : ; : : : , : ‘ : : oe ELS
Trigonopsis Perty . , ; : ; i : 5 ; : : . 120
Keys to species. ‘ ; : ‘ : : : : : A ole
The rufiventris-group. : ; ’ - ; : : , : = 30
The vicina-group . ; : : : : ; ; : : : 2 39
The intermedia-group . : , : ; : : ; : : . 143
The neotropica-group . , ; ; ; : c F ‘ ; ead
References ' : ‘ , ; i : ; , ; : 2 Selo
Index . ‘ : : : 2 ; F ‘ ‘ : ; : rma bP
Synopsis
Keys are provided to the sixteen species of Trigonopsis Perty here recognized, of which eight are described
as new. Two specific names are newly placed in synonymy. The species are placed in four species-groups,
one of which consists entirely of newly described species. Previously published information on biology is
summarized, and further biological observations are added where possible.
Introduction
The wasps of the genus Trigonopsis are uncommonly encountered inhabitants of the New World
tropical forests. They are strikingly elegant creatures, very elongate with a triangular, prognathous
head, petiolate gaster, and most parts highly polished. The mandibles are also long and slender,
well-adapted to catching and carrying the cockroach prey, which are stored in mud nests. These
wasps are scarce in collections, since most collectors have tended to avoid forest habitats with their
relatively low yields of insects.
Several species were described by various early authors, notably Smith (1851; 1856; 1873),
usually on the basis of single specimens. Kohl (1902) first revised the genus, treating it as a sub-
genus of Podium. Since he did not see Smith’s types, misunderstood the descriptions, and in most
cases saw no other material assignable to Smith’s species, he placed in synonymy several names
since regarded as valid. He also confused other species and was misled by inaccurately labelled
specimens.
Richards’ (1937) revision was more limited in scope, since its main purpose was to provide
names for the specimens collected during an expedition to Guyana. However, it also included
the material already in the collections of the British Museum (Natural History), where most of
Smith’s types are housed. Richards, combining this advantage with a more critical approach,
clarified much of the previous confusion.
Menke in Bohart & Menke (1976), in the context of a revision of sphecid genera, divided
Trigonopsis into the rufiventris and intermedia species-groups, and stated that the latter is the
more generalized of the two. However, the present study attempts to show that the intermedia-
group as defined by Menke is more conveniently treated as three distinct groups. Nomenclature
in the genus was also updated.
Bull. Br. Mus. nat. Hist. (Ent.) 37 (4): 117-152 Issued 27 July 1978
117
Examination of recently acquired material in the British Museum (Natural History), using
Richards’ (1937) paper, indicated that undescribed species were represented. It also became evident
that some characters, especially those of the female mandibles and male genitalia, had never
received sufficient attention from any author. These facts together stimulated the present work.
Due mainly to lack of material, no previous author was in a position to assess the proximity
of species and the extreme variation within some of them. In particular, the absence of the opposite
sexes of many species frequently gave rise to misassociations of those present, and to other
misidentifications. Considerable efforts were therefore made to gather together all available
material, and a very wide range of characters was assessed. In addition, several sex associations
were confirmed by reared specimens. As a result, a foundation has been laid for the detailed
evaluation of supraspecific relationships here attempted. A high proportion of new species is
described within this framework, with very few sexes remaining unknown. The intermedia-group
of Menke in Bohart & Menke (1976) is here divided into three, including a species-group con-
sisting entirely of newly described species.
Certain body measurements, and some ratios derived from them, proved to be of value in
separating species. Measurements were made on most specimens with the exception of the
numerous specimens of violascens, rufiventris and cameronii, of which samples were subjectively
selected.
All measurements in the keys and descriptions are in millimetres, and ratios are expressed as an
index derived from the division of one measurement by another. They are abbreviated as follows.
BL Body length, excluding appendages. ;
Notch Female mandible always has a notch of varying size and shape in the inner margin or on the
inner side, nearer apex than base.
MBL Mandibular base length. From notch to base of eye.
MAL Mandibular apex length. From notch to tip.
MR Mandibular ratio. MBL divided by MAL. (When the mandible is very worn, the abnormally
high MR resulting is given in brackets after the normal range.)
CTW Clypeal tooth width. In the females the distance between the tips of the outermost clypeal teeth.
(Most males have only 2 teeth, or the clypeus is distinct.) nee
CED _Clypeal emargination depth. Perpendicular distance from the base of the central emargination
to the line joining the tips of the teeth (males only). ;
TOD Tooth to orbit distance. From the tip of a tooth used to measure CTW, to the nearest point of
an orbit.
CR Clypeal ratio. CTW divided by TOD.
CER Clypeal emargination ratio. CTW divided by CED (males only).
HW Head width. Maximum, including eyes. we:
HL Head length. From the base of a tooth nearest centre of clypeus, to posterior edge of occipital
carina.
HR Head ratio. HW divided by HL.
LIW Lower interocular width. Least distance between lower orbits.
MIW_ Middle interocular width. Greatest distance between middle orbits.
UIW _ Upper interocular width. Least distance between upper orbits.
EL Eye length.
PDL _ Propodeal dorsal length.
Po Petiole length. Distance from the apex of the propodeal valve to the junction with tergite 1.
PR Petiole ratio. PL divided by PDL.
Material studied in the present work is deposited in the institutions abbreviated as follows.
AMNH, New York American Museum of Natural History, New York.
ANS, Philadelphia Academy of Natural Sciences, Philadelphia.
BMNH British Museum (Natural History), London.
BPBM, Honolulu Bernice P. Bishop Museum, Honolulu.
CM, Pittsburgh Carnegie Museum, Pittsburgh.
CMNH, Los Angeles County Museum of Natural History, Los Angeles.
CNC, Ottawa Canadian National Collection, Ottawa.
CU, Davis California University, Davis.
118
CU, Ithaca Cornell University, Ithaca, New York.
Fritz coll. Personal collection of M. A. Fritz, Instituto Entomolégico San Miguel, San
Miguel, Province of Buenos Aires.
FSAE, Gembloux Faculté des Sciences Agronomique de l’Etat, Gembloux, Belgium.
IML, Tucuman Instituto Miguel Lillo, Tucuman, Argentina.
IRSNB, Brussels Institut Royal des Sciences Naturelles de Belgique, Brussels.
IZA, Maracay Instituto de Zoologia Agricola, Maracay, Venezuela.
MCZ, Harvard Museum of Comparative Zoology, Harvard.
Menke coll. Personal collection of Dr A. S. Menke, U.S. Dept of Agriculture, c/o US.
National Museum, Washington.
MHN, Geneva Muséum d’Histoire Naturelle, Geneva.
MIZSU, Turin Museo ed Istituto di Zoologia Sistematica dell’ Universita, Turin.
MNHBN, Paris Muséum National d’Histoire Naturelle, Paris
MNHU, Berlin Museum fiir Naturkunde der Humboldt-Universitat, Berlin.
MP, Belém Museu Paraense Emilio Goeldi, Belém, Para, Brazil.
NM, Rotterdam Natuurhistorisch Museum, Rotterdam.
NM, Vienna Naturhistorisches Museum, Vienna.
NR, Stockholm Naturhistoriska Riksmuseum, Stockholm.
RNH, Leiden Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands.
TM, Budapest Természettudomanyi Muzeum, Budapest.
UM, Oxford University Museum, Oxford
USNM, Washington United States National Museum, Washington, D.C.
UZM, Copenhagen University Zoological Museum, Copenhagen.
ZSBS, Munich Zoologische Sammlung des Bayerischen Staates, Munich.
Acknowledgements
I wish to acknowledge the help of: Mr M. Cooper, for making special efforts to collect these
wasps and their nests; Prof. O. W. Richards, for valuable help and encouragement; Mrs J. A.
Marshall and Dr A. B. Gurney for naming cockroaches and crickets. I especially wish to thank
Dr A. S. Menke for making available his personal notes and specimens.
I thank the following for reading the manuscript and making useful suggestions: Mr M.
Cooper, Dr M. J. W. Eberhard, Dr W. G. Eberhard and Dr A. S. Menke.
I am particularly grateful to Mr M. C. Day for his painstaking reading of the manuscript and
for testing the keys.
I also wish to thank the following persons for loans, gifts or sale of specimens: Dr Cl. Besuchet
(MHN, Geneva), Prof. R. M. Bohart (CU, Davis), Dr P. Dessart (IRSNB, Brussels), Dr E. Diller
(ZSBS, Munich), Dr M. J. W. Eberhard (Universidad del Valle, Cali, Colombia), Dr P. P.
d’Entréves (MIZSU, Turin), Dr S. Erlandsson (NR, Stockholm), Prof. H. E. Evans (Colorado
State University), Mrs M. Favreau (AMNH, New York), Dr F. Fernandez Yépez (IZA,
Maracay), Dr M. Fischer (NM, Vienna), Mr M. A. Fritz (Instituto Entomoldgico, San Miguel,
Buenos Aires), Mrs C. N. Higa (BPBM, Honolulu), Dr P. G. Howes (Bruce Museum of Natural
History, History and Art, Greenwich, Connecticut), Dr S. Kelner-Pillault (MNHN, Paris), Dr
E. Konigsmann (MNHU, Berlin), Dr J. Lawrence (MCZ, Harvard), Prof. J. Leclercq (FSAE,
Gembloux), Dr J. P. van Lith (ZM, Amsterdam), Dr O. Lomholdt (UZM, Copenhagen), Dr
L. Masner (CNC, Ottawa), Mr R. C. Miller (CU, Ithaca), Miss G. Nakahashi (BPBM,
Honolulu), Mr C. O’Toole (UM, Oxford), Dr William L. Overal (MP, Belem), Dr J. Papp
(TM, Budapest), Prof. L. L. Pechuman (CU, Ithaca), Dr B. Petersen (UZM, Copenhagen), Dr
D. C. Rentz (ANS, Philadelphia), Dr J. G. Rozen (AMNH, New York), Mr José M. Schunke
(Pucallpa, Peru), Mr R. R. Snelling (CNMH, Los Angeles), Dr P. M. F. Verhoeff (Utrecht,
Netherlands), Margaret K. Thayer (MCZ, Harvard), Dr G. E. Wallace (CM, Pittsburgh), Dr S.
C. Willemstein (RNH, Leiden) and Dr A. Willink (IML, Tucuman).
119
TRIGONOPSIS Perty
Trigonopsis Perty, 1833 : 141, pl. 27, fig. 18. Type-species: Trigonopsis abdominalis Perty, 1833 (= Podium
rufiventre Fabricius, 1804), by monotypy.
Trigonopsis Perty; Smith, 1851 : 31; 1856 : 226, pl. 6, fig. 1; 1873 : 54.
Trigonopsis Perty [as subgenus of Podium Fabricius]; Kohl, 1902 : 2.
Trigonopsis Perty; Williams, 1928 : 114.
Trigonopsis Perty; Richards, 1937 : 107.
Trigonopsis Perty; Menke in Bohart & Menke, 1976 : 96.
The following description is largely based on that given by Menke in Bohart & Menke (1976:
96-97), with certain modifications.
Highly polished wasps 13-26 mm long, black with red gaster or entirely metallic blue-violet. Forewing
hyaline or slightly infuscate with 2 dark, transverse bands towards apex. Head large, + triangular,
strongly prognathous. Occipital carina sometimes expanded into a flange. Mandible long and slender,
especially in females, in which its inner margin always has a notch of varying size and shape near the apex,
and usually also a basal spine. Some males have a sub-basal tooth on the mandibular inner margin.
Female clypeus with 4-7 marginal teeth, male with 2 or structure quite different. Some males have sub-
clypeal tubercles. Antennae cylindrical, sometimes flattened or expanded in males. Pronotum + elongate,
often with a dorsal projection. Mesopleuron with complete episternal sulcus. Propodeum with spiracular
groove, its dorsum (= metapostnotum) with a longitudinal groove containing a carina and strong
transverse rugae. Remainder of dorsum with or without transverse rugae. Gaster with long petiole.
Among the Sphecinae, Trigonopsis is easily distinguished by its elongate shape, prognathous
head and typical colour patterns, especially of the wing-bands. These characters are shared by
only a few of the species of Podium Fabricius and Penepodium Menke, its closest relatives (these
two genera are hereinafter referred to as Podium s.1.). Trigonopsis is distinguished from these
genera by its propodeal dorsal groove containing a carina and transverse rugae (at most a shallow,
simple groove in Podium s.|.), the spiracular groove (absent in Podium s.1.) and the complete
episternal sulcus (incomplete below in Podium s.1.). In addition, the female of Trigonopsis always
has a mandibular notch (although vestigial in one species, it is never absent), and the cuspis of
the male genitalia has a basal post (Fig. 70). Both notch and post are lacking in Podium s.1.
THE SPECIES-GROUPS. Sixteen species are recognized. Since the key is artificial, it does not reflect
the four natural groups in which these are placed. They are characterized and inter-related as
follows.
rufiventris-group vicina-group
violascens (Dalla Torre) richardsi sp. n.
violaceus Smith vicina (Dalla Torre)
resplendens (Kohl) syn. n. affinis Smith
menkei sp. n. schunkei sp. n.
rufiventris (Fabricius)
abdominalis Perty intermedia-group
haemorrhoidalis Smith intermedia Saussure
soror Mocsary syn. n. cyclocephalus Smith
frivaldskyi Mocsary moraballi Richards
cameronii (Kohl)
howesi sp. n. neotropica-group
grylloctonus Richards neotropica sp. n.
succinea sp. n.
mocoana sp. n.
cooperi sp. n.
The rufiventris-group. Propodeal dorsum with, at most, traces of rugae outside central furrow. 3
sternite 8 bifid, genitalia distinctive, basal post of cuspis round-ended. ? clypeus with 5+ regular teeth,
mandible with V-shaped notch. 3 without subclypeal tubercles, antennae cylindrical.
The vicina-group. Propodeal dorsum with rugae outside central furrow. ¢ sternite 8 entire, basal post
of cuspis round-ended. 2 clypeus with 4-5 regular teeth, mandible with U- to V-shaped notch. 3 with
subclypeal tubercles and flattened, + expanded antennae.
120
The intermedia-group. Propodeal dorsum with rugae outside central furrow. ¢ sternite 8 entire, basal
post of cuspis round-ended. 2 clypeus with 6—7 often irregular teeth, mandible with U-shaped notch. 3g
without subclypeal tubercles, with cylindrical antennae.
The neotropica-group. Propodeal dorsum with rugae outside central furrow. ¢ sternite 8 entire, basal
post of cuspis square-ended. 2 clypeus with 5 sometimes irregular teeth, mandible with step-like notch.
$ sometimes with subclypeal tubercles, and usually slightly flattened antennae.
The vicina-group shows certain characters in common with all other groups of the genus. The
clypeal and mandibular shape of female richardsi, the occipital flange of both its sexes, and the
HR of 1:0 of female richardsi and all vicina-group males, are characters otherwise typical of
the rufiventris-group.
The females of vicina and schunkei, the other two species of the vicina-group, have the mandibles
strongly resembling those of all the intermedia-group females. These two groups are further linked
by the males of intermedia and schunkei, the only two in the genus which have a sub-basal tooth
on the mandibular inner margin. They are also the largest species of their respective groups.
The subclypeal tubercles found in all vicina-group males are also present in the neotropica-
group, in the male of neotropica itself. The strong flattening and partial expansion of the antennae
in all vicina-group males is paralleled by the slight flattening found in all known neotropica-group
males except that of cooperi. The unusually dense pubescence of the anterior mesoscutum is
found in several species of both groups, but not usually in both sexes of any one species. The first
two characters are also found in certain species of Podium and Penepodium, which may indicate
that they are ancestral.
DIsTRIBUTION. Most species of this genus are confined to lowland forest, a few ascending to be-
tween 600 and 1000 m. The range extends from Mexico to west-coast Ecuador, and everywhere
east of the Andes south to Bolivia, the Mato Grosso and Rio de Janeiro.
VARIATION. The back of the head is sometimes greatly expanded, at times almost quadrate. This
appears to be purely individual variation and is found in all species-groups. It occurs mainly in
larger specimens and is more common in males than in females.
In addition, various structures show variation along two clines. From the east, one can be
traced north-west into the Guianas, the other west along the Amazon. The latter is also accom-
panied by a colour cline. The Guianas are relatively isolated by high mountains to the south and
west. Several species are so far known only from this area. The Andes form the western limit of
distribution for all the Amazon Basin species.
The structural clines are most marked in violascens, neotropica and cooperi; details are given
under those species.
Colour variation is most easily observed in rufiventris as it is the most widespread and commonly
collected species of the genus. Almost all the specimens from eastern Brazil and the Guianas are
very dark, becoming paler on average westwards along the Amazon. The red colour changes to
orange and becomes more extensive, and the wings assume a strong amber tint. West of about
68° W., extreme, pale specimens occur (= soror Mocsary). All the females from Chanchamayo,
Perené, Tapiche, Pebas, Iquitos and Sao Paulo de Olivenga belong to this form. It is interesting
that it seems to occur in isolation, since less extreme forms are unknown from these localities.
There are reasonably long series from Mocoa and Pucallpa which show a good range of colour
variation, but do not include the extreme form. The phenomenon affects other species in different
localities. For example, a female richardsi from Satipo represents the extreme, pale form, while a
female rufiventris from the same locality does not.
The females of all the red-and-black species show the same general tendency to become paler
westwards along the Amazon, but to differing degrees. In cooperi, for example, the tendency is
already strongly evident at 55° W., but not until further west in other species. Although males
follow the same general trend, they are darker on average, and extreme, pale forms are unknown.
BioLoGy. Eberhard (1972) noted interactions between individuals of cameronii while they were
provisioning nests. This is of particular interest in that the behaviour may be described as sub-
social. With reference to the same species, Eberhard (1974) gave the most comprehensive account
of biology in the genus, including comparisons with certain species of other genera.
121
So far as is known, all species of Trigonopsis prey on cockroaches (apparently usually nymphs)
except for gry//octonus and rufiventris, reported to prey on crickets (Richards, 1937 : 111, and the
present study, p. 134). The prey are stored in cells made of mud, several in each cell. Nests con-
sist of one to many cells and are attached to a great variety of forest substrates above ground
level, of various degrees of stability. Usually only small nests are found under leaves, so that
larger species tend not to choose such places. However, when smaller species choose firmer sites
such as rock-faces, they usually make much larger nests than otherwise. On rock-face and tree-
trunk sites, large numbers of cells often agglomerate and old ones are frequently renovated for
re-use. Variation in the surface structure of mud added after the cells are completed may possibly
serve the purpose of camouflage. The cell axes are always parallel and normally horizontal, those
recorded by Richards (1937 : 109-110) being exceptional. Up to 4 cells may be placed ina single
row but they form a bundle if more numerous. Further collecting is desirable to establish how the
size and structure of nests varies according to species, site and region. At least some species
belonging to different groups have differently shaped cocoons (Figs 36-39).
FUNCTIONAL MORPHOLOGY. Eberhard (1974 : 304307) has already suggested how the clypeus and
mandibles interact when cameronii catches and carries its prey. The sharp sides of the cockroach
prey are well gripped during capture by the mandibular notches of the female wasp. Subsequently,
these notches oppose the clypeal tooth gaps to grip the palpi of the prey, and prevent it slipping
forwards or backwards between the wasp’s legs, which hold it during the flight to the nest. A
comparison of different species and species-groups reveals interesting compensations. The clypeal
teeth of the intermedia-group species are usually very blunt and irregular, but the mandibles have
broad notches and are the stoutest in the genus. However, most species of the rufiventris-group
have very long, sharp, clypeal teeth, and mandibles with a strong expansion before a narrow notch,
whose effect is further enhanced by the sharp bend of the mandible around it. The clypeal teeth
of violascens are more variable and often less sharp, but the mandibular expansion is distinctly
angulate. Related genera, which never have a mandibular notch, solve the problem of grip in two
ways. Podium females have slender mandibles but many clypeal teeth, while those of Penepodium
usually have stouter mandibles, which are angulate below, and only 2 clypeal teeth. (See also
rufiventris-group notes.)
Key to species
It is important to open the mandibles, especially of females. Mandibular expansions are best observed
dorsolaterally.
Females
The female of grylloctonus is unknown.
1 Body entirely dark, with violet-blue metallic lustre . ‘ ; : : . 2
— Head and thorax black, gaster except petiole red (sometimes dark red) é 3
2 Pronotal dorsum in profile shallowly, evenly convex. Petiole of even width (Fig. 1). (West of
the Andes, north to Mexico) . : cameronii (Kohl) (p. 136)
— Pronotal dorsum in profile with weak, angular projection posteriorly (Figs 16, 17). Petiole
slightly expanded posteriorly (Figs 2-4, 6-8, 10). (Widespread east of the Andes)
violascens (Dalla ee a 131)
3 Pronotal dorsum ‘in profile shallowly, evenly convex ‘ ; : 4
— Pronotal dorsum in profile with sharply angulate projection posteriorly : ' 6
4 Petiole with small but distinct expansion just before junction with tergite 1 (Figs 9, 11)
menkei sp. n. (p. i:
Petiole of even width .
5 Pronotal dorsum with shallow but distinct longitudinal groove : $ howesi sp. n. (p. ian
— Pronotal dorsum with at most an indistinct longitudinal groove rufiventris (Fabricius) (p. 133)
6 Clypeus with 6 or more teeth, outermost pair strongly bent forward, those between often
weak or irregularly spaced. Mandible stout, evenly tapered, with large notch (Figs 12-15,
22,.25)-.% : ; : : , . ; ; ; : ; : ; E ¥
122
rufiventris Q
violascens Q merce: )
1
ic
moraballi
$ intermedia Q
violascens Q
Figs 1-17 Trigonopsis spp. (1-5) heads and clypeal variants. (6-11) petioles. (6-9) left side.
(10, 11) dorsal view. (12-15) heads and clypeal variants. (16, 17) pronota, left side.
123
10
11
[2
13
14
Clypeus with 4 or 5 teeth, usually well formed and regularly spaced, outermost pair frequently
weakly bent forward or straight. If mandible stout, inner side at least slightly expanded
before notch, which is often small 9
HW at least 4-4. BL 22-27. Mandibular inner margin without basal spine. “CTW at least He 5
(Pigs 145:15)/.- : , intermedia Saussure (p. 143)
HW at most 4-5. BL 13-22. A small, basal spine or tooth present on mandibular inner margin.
CTW at most 1:5. 8
4 smaller teeth of clypeus fairly equal i in size and spacing. HW 4: 1-4- 2, Mandibular notch not
overhung by a carina, apical part of mandible much more slender than basal (Figs 12, 13)
moraballi Richards (p. 145)
4 smaller teeth of clypeus often irregular, centre pair separated by a wide, shallow gap which is
sometimes filled by a blunt, vestigial seventh tooth. HW 4:3-4-5. Mandibular notch partly
overhung by a strong carina, apical part of mandible evenly tapered with basal (Figs 22, 23)
cyclocephalus Smith (p. 144)
Outermost pair of clypeal teeth equal to or slightly shorter than adjacent pair. Mandible
with notch small and step-like, before which the inner margin is moderately ica
(Figs 18-20, 24, 25) . ; 10
Outermost pair of clypeal teeth distinctly longer than adjacent pair. "Mandible often not as
above. 11
BL 22-26. CR i 8-2: 2, Pe oy 144, Propodeal dorsum ‘with | coarse rugae outside central
groove, which is deep throughout. Small, basal spine on mandibular inner margin rather
broad, rounded at tip. Top of pronotal projection very strongly raised, in profile with
distinct concavity in front (Fig. 28). Wings strongly amber-tinted, stigma pale. Clypeus
with 2 central gaps between teeth deeper than outer ones. HW 3-8-4-6 (Figs 24, 25)
succinea sp. n. (p. 149)
BL 19-20. CR 2:7-2:9. PL 2-9-3-2. Propodeal dorsum with fine rugae outside central groove,
which is shallow anteriorly. Small basal spine on mandibular inner margin narrower,
sharply pointed. Top of pronotal projection less strongly raised, scarcely concave in front
(Fig. 27). Wings not amber-tinted, stigma dark brown, most veins dark. Clypeus with 2
central gaps equal to or shallower than outer ones. HW 3-7-3-9 (Figs 18-20)
neotropica sp. n. (p. 147)
Antennal segment 3 longer than 4 and 5 together. Mandible slender, evenly tapered, rather
strongly curved, with distinct, U- to V-shaped notch. MR 1-9-3:2. Clypeus with 2 ah
teeth and 3 smaller ones between (Figs 21, 26) . 12
Antennal segment 3 not longer than 4 and 4 together. Mandible variously shaped but. not as
above, notch sometimes small or vestigial. MR at least 2-4. Clypeus not always as above . 13
MR 2:7-3-2 (-5:6) MIW 2:2-2:4 times CTW. Antennal segment 3 is 2:5—2:8 times as long as 6.
CR 1-3-1-6. Centre tooth of clypeus equal to or slightly ie than adjacent pair. BL 16-19.
HW 3:-1-3-7 (Fig. 21) , vicina (Dalla Torre) (p. 141)
MR 1-9-2:3. MIW 1-7-2:1 times CTW. Antennal segment 3 is 2:9-3-3 times as long as 6.
CR 1-6-2:1. Centre tooth of clypeus equal to or shorter than adjacent pair. BL 18-22.
HW 3-7-4-4 (Fig. 26) : : schunkei sp. n. (p. 142)
HW 2:8-3-5, at most 0-1 more than HL. Occipital carina + expanded into a thin, lamellar
flange, often translucent. BL 13-17. Mandibular inner margin + strongly expanded before
V-shaped notch. Mandible usually sharply bent at notch and abruptly tapered after it, upper
edge of apex inflexed near notch (Figs 29-32) . ; : richardsi sp. n. (p. 140)
HW 3-5-4-0, at least 0-66 more than HL. Occipital carina very strong but not forming a flange.
BL 17-20. Mandibular inner margin at most moderately expanded before step-like notch.
Mandible often shallowly bent at notch, rarely abruptly tapered after it. Upper edge of
apex not inflexed . ; 14
Mandible stout, inner margin moderately expanded before notch, which i is small but distinct.
MR 3-4-3-8. EL 2:2-2:3. Toothbearing area of clypeus more projecting, margin outside
teeth at steep angle, so that CR is 1-3-1-7; all clypeal teeth bent forward, outermost pair
most strongly (Fig. 35). mocoana sp. n. (p. 149)
Mandible slender, evenly tapered, notch vestigial. ‘MR 4: 6-5: 5 (-7: 3), EL 2-0-2:3. Tooth-
bearing area of clypeus less projecting, margin outside teeth at shallower angle (where almost
sympatric with mocoana, forming a broad, translucent lamella), so that CR is 1:8-2-7;
only outermost pair of teeth bent forward, contrasting with others (Figs 33, 34)
cooperi sp. n. (p. 150)
124
neotropica fe)
SAS schunkei 2
ry 27 neotropica fe)
\
meee
28 succinea Q
——
richardsi 2
succinea Q
30
richardsi 9
richardsi Q richardsi fe)
Figs 18-32 Trigonopsis spp. (18-26) heads and clypeal variants. (27, 28) pronotal dorsa, left side.
(29-32) heads.
125
OF,
eb
mocoana © violascens 39
4
violascens &!
J Pe ieee 3
ee AT 48 ee
violascens Bo" violascens 6'o! 50 grylloctonus 3
Figs 33-51 Trigonopsis spp. (33-35) heads and clypeal variants. (36-39) cocoons. (40-42) heads.
(43-50) petioles showing (43-46) left side; (47-49) dorsal view; (50) left side. (51) head.
126
92
succinea
schunkei &
schunkei 3
richardsi 3
richardsi 3
moraballi 3
cooperi 3!
intermedia & 4 y
Figs 52-65 Trigonopsis spp. (52-55) heads. (56-59) antennae. (60-65) heads and clypeal variants.
127
66 cyclocephalus 3
is
71 73 75
68 cooperi TOrcomerchicn he Sens So Téa howe! LO eas F
69
67. moraballi 7
.]} BASAL POST
4 OF CUSPIS
if
78 Sree ws of 80 schunkei 3 82 richardsi 3 84 vicina S' 86 neotropica rot
Figs 66-86 Trigonopsis spp. (66-68) sternites 8. (69-86) aedeagi and volsellae.
128
Males
The males of menkei and mocoana are unknown.
1
2
13
Body entirely dark, with violet-blue reflections
Head and thorax black, gaster except petiole red (sometimes dark red) ;
Clypeus with 2 large, sharp teeth (Fig. 40). Pronotal dorsum in profile shallowly, evenly
convex. Petiole in dorsal view of uniform width throughout. Genitalia as in Figs 69, 70.
(West of the Andes, north to Mexico) : cameronii (Kohl) (p.
Clypeus with large, central truncate projection (Fig. 41), Pronotal dorsum in profile, with
weakly angular posterior projection (cf. Figs 16, 17). Petiole in dorsal view, strongly
expanded at junction with tergite 1 (Figs arcs Genitalia as in Figs 71, 72. (Widespread
east of the Andes) . ; violascens (Dalla Torre) (p.
Pronotal dorsum in profile, shallowly, evenly « convex
Pronotal dorsum in profile, with sharply angulate projection posteriorly
Petiole strongly expanded in middle (Figs 49-51). Genitalia as in Figs 75, 76
grylloctonus Richards (p.
Petiole of even width throughout .
Pronotal dorsum with shallow longitudinal groove. Genitalia as in Figs (bx "14 howesi sp. n. (p.
Pronotal dorsum without any trace of a longitudinal groove (Fig. 42). Genitalia as in Figs 77,
78. F rufiventris (Fabricius) (p.
Clypeus with 2 large, rectangular, ‘divergent lobes. Mandibular j inner margin with sub-basal
tooth (Fig. 52).
Antennae as in Fig. 56. Genitalia as in Figs 79-80 . ; schunkei sp. n. (p.
Clypeus with 2 large, sharp teeth, its tooth-to-orbit margin with at most a notch. Mandibular
inner margin sometimes with sub-basal tooth
Antennal segments moderately to strongly dorsoventrally flattened, their junctions offset in
lateral view (Fig. 59). Mandibular inner margin without sub-basal tooth
Antennal segments almost perfectly cylindrical, their junctions not or scarcely offset in lateral
view. Mandibular inner margin sometimes with sub-basal tooth
Clypeal tooth-to-orbit margin entire and without any tubercles behind (view from below)
(Fig. 53). BL 17-20, HW 3-3-3-8, HL 3-0-3-5, PL 2-7-3°8.
Genitalia as in Figs 95,96. ‘ succinea sp. n. (p.
Clypeal tooth-to-orbit margin with at least 2 tubercles behind (view from below), sometimes
also with a deep notch visible in front view. BL 13-18, HW 2:7-3-6, HL 2:7-3-3, PL 2:3-2:9
Clypeal tooth-to-orbit margin in front view, with a deep U-shaped notch. Seen from below,
sides of notch are tuberculate behind, and outer tubercle is connected to a third one still
further back (Fig. 55). Antennal segments 5 and 6 slightly ioe tae (Fig. 57).
Genitalia as in Figs 81,82 . : : richardsi sp. n. (p.
Clypeal tooth-to-orbit margin almost entire in ‘front view but seen from below, base of outer
side of each tooth is tuberculate behind. Beyond it, and behind tooth-to-orbit margin but
not forming part of it, is another tubercle. The two tubercles together form a deep notch,
and tip of second is just visible in front view. Antenna not as above (Figs 54, 60) :
Antennal segments 5-10 strongly expanded (Fig. 58). Eyes strongly divergent downwards
(Fig. 60). Genitalia as in Figs 83, 84 . : : : vicina (Dalla Torre) (p.
No antennal segments expanded, only flattened. Eyes scarcely divergent downwards (Fig. 54).
Genitalia as in Figs 85, 86 ; ; neotropica sp. n. (p.
Mandibular inner margin with sub- basal tooth. BL 19-5- 23: aU HW 3-9-4:5, CTW 0:90-1:05,
CDR 1-4-1-7, CR 0-9-1:1 (Figs 62, 63)
Genitalia as in Figs 87,88 . : intermedia Saussure (p.
Mandibular inner margin without sub-basal tooth. BL 15: 0-19: 0, HW 3:-1-3-7, CTW 0:30-0:40,
CDR 0-9-1:1,CR0-40_.
TOD 0:85-0:97, PDL 1-8-2:2, third antennal segment ihe i= 3 long and 1: g- 1-9 times a as long
as the first, eyes slightly divergent below, CED 0-35-0:40, HL 2-8-3-2, HW 3:-1-3-7,
CTW 0-35-0-40, PL 2:3-3-0 (Fig. 63) (Upper Amazon).
Sternite 8 as in Fig. 66. Genitalia as in Figs 89,90 . ; cyclocephalus Smith (p.
TOD 0:70-0:77, PDL 1-7-1-9, antennal segment 3 is 0-9-1-0 long and 1-5—1-6 times as long as
the first, eyes parallel or slightly convergent below, CED 0:27-0:32, HL 2:8-2-9, HW 3-1-3°3,
CTW 0:30-0:35, PL 2:4-2:6. (Amazon and Guyana) .
MIW 0-35 more than UIW, PDL 1-7-1-8, PL 2:4-2:5, CTW 0- 35, CED 0- 32 (Fig. 61). Sternite
129
WN
136)
149)
140)
10
141)
147)
143)
12
145)
13
8 rather sharply keeled, its tip broadly rounded (Fig. 67). Genitalia as in Figs 91, 92
(Guyana) ; f ; , , ; : : : . moraballi Richards (p. 146)
MIW 0:15-0:20 more than UIW, PDL 1-8-1:9, PL 2:6, CTW 0:30-0:32, CED 0:27-0:30
(Fig. 65). Sternite 8 obtusely keeled, its tip narrowly truncate (Fig. 68). Genitalia as in
Figs 93, 94. (Amazon mainstream from Tena to Piaui, and East Peru) . cooperi sp. n. (p. 150)
N.B. The last three males are extremely similar externally, although cooperi is easily dis-
tinguished by its genitalia. The differences between the genitalia of cyclocephalus and moraballi,
although slight, appear to be constant.
cooperi &
succinea &
92 raotabale of 95
90 cyclocephalus 3
Figs 87-96 Trigonopsis spp., aedeagi and volsellae.
88 intermedia 3
The rufiventris-group
Unique characters are: complete or almost complete lack of transverse rugae outside the central furrow of
the propodeal dorsum in both sexes, and bifid sternite 8 and distinctive genitalia of the males. Further
characters are: small, slender species with HR of 1-0 or less, very broad occipital flange. Female mandible
sharply bent, inner margin with V-shaped notch and strong expansion before it, 5 usually sharp clypeal
teeth, sternite 6 sharply keeled. Cuspis of male genitalia with round-ended basal post. Dark parts, at least
of legs, + metallic in all species.
Within the group, violascens is distinguished by the dorsal pronotal projection, lacking in the
other species. Its male is further distinguished by the unique shape of its clypeus and petiole,
and has the most distinct genitalic structure in the group. The female mandibular expansion,
rounded in the other species, is angulate and the clypeal variation considerably greater.
This group is by far the most common and widespread of all. It contains the only species found
west of the Andes. It includes all the known metallic species, and it is interesting that both of the
entirely metallic-coloured species have a greater altitude range than any others of the genus. The
130
group also includes the only species which lack a pronotal projection, and the single species known
to have diverged in its prey choice. Furthermore, it appears that the group may recently have
evolved rapidly, since with the exception of violascens, the species are very closely related (for
example, the head ratios of the species of this group are much more uniform than those of the
species of other groups). They are usually separated by small and often single structural characters,
although sometimes also by considerable biological ones.
The species rufiventris, cameronii and violascens are apparently more common than any other
Trigonopsis. Various factors may contribute to their frequency, such as: the unusual shape of the
female mandibles and clypeus affording better grip of the prey; small average size reducing the
need to find large or numerous prey; greater adaptability in choice of nesting site. Certainly
richardsi, much the commonest species of the vicina-group, is very similar in at least the first two
respects (its biology is unknown). The remaining species of the rufiventris-group, all of them
apparently much less widely distributed, may represent specialized adaptations to certain very
localized ecological niches.
Trigonopsis violascens (Dalla Torre)
(Figs 2-4, 6-8, 10, 16, 17, 36, 37, 41, 43-48, 71, 72, 99)
Trigonopsis violaceus Smith, 1851: 31. Holotype 3, BRAzit (BMNH) [examined]. [Junior secondary
homonym in Sceliphron Klug, of Sphex violacea F., 1775.]
Sceliphron violascens Dalla Torre, 1897 : 392. [Replacement name for Trigonopsis violaceus Smith.]
[Podium (Trigonopsis) cameronii Kohl, 1902 : 39, 2 (part). Misidentification.]
Podium (Trigonopsis) violaceum Smith; Kohl, 1902 : 39, pl. 1, fig. 3, pl. 5, figs 55, 63 [not pl. 6, fig. 63 as
stated in text], pl. 6, figs 70, 80, 3.
Podium (Trigonopsis) resplendens Kohl, 1902 : 41, pl. 1, fig. 1, pl. 5, fig. 61, pl. 6, figs 72, 74 [actually an
illustration of violaceum], pl. 7, fig. 85. Holotype 2, BRAzIL (NM, Vienna) [examined]. [Synonymy by
Richards, 1937 : 109.]
Trigonopsis violaceus Smith; Richards, 1937 : 109, 112.
Trigonopsis resplendens Kohl; Menke in Bohart & Menke, 1976: 98.
Trigonopsis violascens (Dalla Torre); Menke in Bohart & Menke, 1976 : 98.
2. BL 15-0-19-5 (but see below). Almost entirely dark, with blue-violet reflections, especially on head and
gaster. Mandibular base stout, with angulate expansion on inner margin before the notch (Fig. 2). Even
in tiny specimens, this angle is still represented by a point. Notch V-shaped, deep and narrow, proximally
obtuse-angled, distally right-angled. Apex more slender than base, with upper edge strongly inflexed,
especially near the notch. Pronotum with weakly angulate dorsal projection, of a shape different from that
found in other species-groups (Figs 16, 17). Petiole slightly expanded at junction with gaster (Fig. 10).
HW 2:8-3-4, HL 2:9-3-4, HR 0-9-1-0, EL 1-9-2:3, UIW 1-4-1-:7, MIW 1-5-1-9, LIW 1-2-1-5, CTW 0-6-0°8,
TOD 0-4-0:6, CR 1:1-1-6, MBL 1-3-1:9, MAL 0-4-0:8, MR 1-8-3-0, PDL 1-8-2:3, PL 2:6-3-3, PR 1-1-1°5.
An exceptionally small specimen from the Mato Grosso differs as follows: BL 10-5, HW 2-0, HL 1°9,
EL 1-4, UIW 1-0, MIW 1-1, LIW 0-8, CTW 0-4, TOD 0-3, MBL 1-0, MR 3-5, PDL 1-0, PL 1:6.
$. BL 15-0—22:0. Inner margin of mandible always with middle and pre-apical expansions, the latter
stronger (Fig. 41). Clypeus with a large, narrowly truncate projection (Fig. 41). (Because of its unusual
shape it was not measured.) Pronotum with a very weakly angular projection (cf. Fig. 16). Petiole strongly
expanded at junction with gaster (Figs 47, 48). Genitalia Figs 71, 72. HW 2-6-3-8, EL 1-6—2-2, UIW 1-3-
2:1, MIW 1-4-2-3, LIW 1-2-2:1, PDL 1-6—2:6, PL 2:8-3-7, PR 1-4~-1-8. Otherwise as 2.
This species is like cameronii in colour, but is nowhere sympatric with it.
VARIATION. The truncated end of the male clypeal projection is often + emarginate. In the female
clypeus, the centre tooth varies from well-developed (about half the length of the outermost) to
vestigial. The adjacent teeth vary considerably in length, from intermediate between the centre and
outermost, to much longer than the outermost. As the latter extreme is approached, the outer
pair of gaps becomes much shallower so that the clypeus takes the form of two bifid projections
with a small tooth between, and also the mandibular expansion becomes more sharply anguiate
131
(Figs 2-4). The projection on the pronotal dorsum, more broadly based than that found in other
species-groups, is normally low. On average, it is stronger in the Guianas (Fig. 16) and still more
so towards the Andean foothills (Fig. 17). The depth of the cleft in the apex of the projection also
varies, but in a less clear pattern. The normally medium-sized punctures of the mesoscutum are
occasionally very fine. The gastral petiole, normally rather strongly upcurved distally, is sometimes
extremely strongly curved, when it also appears thicker than usual. Ventrally, it has 0-2 rather
strong angles in males (Figs 43-46) and sometimes a single weak one in females (Figs 6-8). In a
male from Leticia the apex of sternite 8 has a single, central spine instead of being bifid. In all
this variation it has not so far been possible to find any consistently coinciding aspects which
could indicate the existence of more than a single species.
DISTRIBUTION. Very widespread east of the Andes, ascending to about 850 m, higher than any
other species in that area (map, Fig. 99). This and neotropica are the only species so far recorded
from the Mato Grosso (see also discussion under cameronii distribution).
BioLoGy. Cooper found the following nests. Colombia: Putumayo, Alto Afan near Mocoa,
19.vi.1974: nest with smooth surface, on rock face. Wasp sitting on nest with four empty cells.
Indistinguishable from rufiventris nest nearby. Amazonas, Leticia, 20.viii.1974: three large nests
attached to rootlets of fallen tree. Two nests (preserved in BMNH) measure about 65 mm high,
42 mm along the cells and 30 mm across them with about 9 cells, and 60 x 27 x 30 mm with about
three cells. The overall shape is rather like that of a hanging pear, with an upper mass of mud
acting as a holdfast, and the cells fixed horizontally below it. The extra mud covering the cells is
rough, solid and irregular, overlaid by very many loose pellets. Cockroach prey taken from one
of these nests consisted of Blattellidae: 2 nymphs of 1 sp., gen. indet. (det. Gurney). Mocoa,
23.iii.1974, suspended from a rootlet under a bank. It had about five cells and (possibly due to
age) a smooth surface. The considerable difference in the form of the final nest-covering used
between the two kinds of nesting-site (rock-face and rootlet) would be consistent with its serving
the purpose of camouflage.
The cocoons were narrow, black, with two bands of white fibrous material (Figs 36, 37).
Richards (1937 : 109) refers to a Dipterous parasite Amobia (as Pachyophthalmus) floridensis
antillarum Richards, bred from a nest of this species in Trinidad.
MATERIAL EXAMINED. Dates and collectors’ names omitted.
Trigonopsis violaceus Smith, holotype 3, BRAZIL: Pard, Belém (Wallace & Bates) (BMNH).
Podium (Trigonopsis) resplendens Kohl, holotype 9, BRAZIL: no further data (NM, Vienna).
FRENCH GUIANA: | 3, Tollinche, River Maroni; 1 9, no further data (MNHN, Paris); 1 9, no
further data (NM, Vienna) (paralectotype of cameronii); 1 9, no further data (MHN, Geneva).
SURINAM: 2 9, | 3, Malaise trap (RNH, Leiden). Guyana: | 9, Bartica (ANS, Philadelphia); 1 9,
Essequibo River, Moraballi Creek (c. 30 km SE. Bartica) (BMNH). Trinipap: | 9, St Augustine
(USNM, Washington); 1 9, 9km N. Arima, 440 m; 3 9, no further data (BMNH). COLOMBIA:
1 9, 1 J, Amazonas, Leticia; 1 9, Amazonas, R. Igara Parana, La Chorrera (mislabelled ‘Peru,
Putumayo District’) (CU, Ithaca); 2 9, 1 3, Putumayo, Mocoa; 2 9, Putumayo, Alto Afan, 7 km
NE. Mocoa; 3 9, 3 3, Meta, Cord. Macarena (BMNH); | 9, further data illegible (UM, Oxford).
PERU: 2 9, Hudnuco, Tingo Maria (BMNH); 1 9, same locality (Menke coll.); 1 9, same locality
(Fritz coll.); 19, same locality, 620m (MCZ, Harvard); 19, Hudnuco, Tingo Maria (Rio
Huallaga), 700m (IML, Tucumdn); 19, Hudnuco, Cord. Azul, Previsto, 850m (BMNH);
1 9, Hudnuco, Pachitea (TM, Budapest); 1 g, Hudnuco, Monzon (Menke coll.); 1 2, Cuzco,
Rio Cosiipata Valley, Hacienda Maria, 400m (IML, Tucumdan); 1 9, Junin, Chanchamayo
(USNM, Washington); 19, 14, same data (Menke coll.); 19, Amazonas, Rio Santiago
(AMNH, New York). Botivia: 1 9, La Paz, Coroico (c. 500m) (MNHU, Berlin). BRAZIL:
2 2, Mato Grosso, 12.50 S., 51.47 W., one reared iii.1968 from mud cells, side of shallow pit; the
other on mud pit, cerradao, 11.ii.1968 (BMNH); 1 9, Para, Belém (RNH, Leiden); 1 9, Para,
R. Arrayolos (MP, Belém); 1 9, Amazonas, R. Japura (MP, Belém); 1 3, no further data (UM,
Oxford); 2 3, no data (USNM, Washington); 1 3g, no data (NM, Vienna).
132
Trigonopsis menkei sp. n.
(Figs 9, 11, 99)
2. BL 22-24. Pronotum without dorsal projection. Gaster red. Petiole with small but distinct lateral
expansion just before junction with gaster (Figs 9, 11). HW 3-8-4-1, HL 3-9-4-2, HR 0-9-1-0, EL 2-6-2:7,
UIW 1-7-1-8, MIW 2:0-2:2, LIW 1-7-1:9, CTW 0-8-1:0, TOD 0-7-0-85, CR 1-1-1:2, MBL 2-0-2°1,
MAL 0:8-1-4, MR 1-5-2-3, PDL 2-7-2:9, PL 3-7-3-9, PR 1:3-1-4. Otherwise as rufiventris.
$. Unknown.
This species differs from its close relatives only in petiole shape.
VARIATION. Mainly in clypeus, as in rufiventris.
DISTRIBUTION. Iquitos and near Tena only (map, Fig. 99).
BioLocGy. Unknown.
MATERIAL EXAMINED. Holotype 2, ECUADOR: Napo, Rio Jatin Yacu (1.04 S., 77.48 W.), 700 m
(MacIntyre) (MCZ, Harvard).
Paratypes. ECUADOR: | 9, same data as holotype (BMNH). Peru: 1 9, Loreto, Iquitos (Bassler)
(AMNH, New York); 1 9, Loreto, Iquitos, 4.viii.1906 (Ducke) (MP, Belém).
Trigonopsis rufiventris (Fabricius)
(Figs 5, 42, 77, 78, 97)
Podium rufiventre Fabricius, 1804 : 184. Holotype 2, SourH AMERICA (UZM, Copenhagen) [examined].
Trigonopsis abdominalis Perty, 1833 : 142, pl. 27, fig. 18. Holotype 2, SourH AMERICA (ZSBS, Munich)
[examined]. [Synonymy by Smith, 1856 : 226.]
Trigonopsis haemorrhoidalis Smith, 1856 : 226, 479. [Unavailable name, inadvertently published in the
synonymy of Podium rufiventre F.]
Trigonopsis soror Mocsary, 1883 : 23. Holotype 2, BRAzIL (TM, Budapest) [examined]. [Synonymy by
Kohl, 1902 : 37.]
Trigonopsis frivaldskyi Mocsary, 1883 : 23. Holotype 2, BRAzIL (TM, Budapest) [examined]. [Synonymy
by Kohl, 1902 : 37.]
Trigonopsis rufiventris (Fabricius); Menke in Bohart & Menke, 1976: 98.
Trigonopsis soror Mocsary; Menke in Bohart & Menke, 1976: 98.
9. BL 15-23. Occipital carina flange-like. Pronotal dorsum without projection and with at most an
indistinct longitudinal groove. Gaster red. Petiole of even width. HW 2-7-3-9, HL 2-7-4-2, HR 0:9-1:0,
EL 1-9-2:6, UIW 1-3-1-7, MIW 1-5-2:0, LIW 1-2-1-:7, CTW 0:65-0:85, TOD 0:50-0:75, CR 1-1-1-6,
MBL 1:35-1:90, MAL 0-7-1-3, MR 1-5-1-9 (—2:-1), PDL 1-8-2:9, PL 2:5-4-0, PR 1-3-1-6. (Fig. 5.)
3. BL 13-5-19-0. Clypeus with two large, sharp teeth. Pronotum with at most the slightest trace of a
longitudinal -groove. Genitalia as in Figs 77, 78. HW 2-6-3:3, HL 2-6-3°5, HR 0-9-1-0, EL 1-8-2:°3,
UIW 1-3-1-7, MIW 1-4-1-8, LIW 1-2-1-6, CTW 0:25-0:32, CED 0:22-0:35, CER 0-9-1-3, TOD 0:57-0°75,
CR 0-4-0°5, PDL 1-6—2:4, PL 2:7-3-7, PR 1-3-1:7. (Fig. 42.) Otherwise as 9.
This species differs from cameronii in colour and distribution, and from other close relatives
in its unmodified pronotum and petiole.
VARIATION. The females show much variation. The five clypeal teeth and the gaps between them
may be subequal; sometimes the outer two pairs of teeth are longer and the centre one reduced
(Fig. 5). The tooth-to-orbit margin is normally straight, but occasionally may be strongly convex
near the tooth. The propodeal dorsal and posterior faces normally form an obtuse angle at their
junction, but occasionally merge in a shallow curve. The propodeal dorsum of large specimens
occasionally bears traces of transverse rugae also outside the central furrow. Only three males of
the seventeen examined had the inner mandibular margin with a preapical expansion, and none
had a middle expansion (Fig. 42). (Cf. violascens Fig. 41.)
(For details of colour variation see general discussion p. 121.)
DisTRIBUTION. Widespread east of the Andes, from Colombia to the Guianas in the north, and
south to Peru in the west and Rio de Janeiro in the east (map, Fig. 97).
£33
rufiventris
extreme, pale
form e
<A
Fig. 97 Collection localities of 7. rufiventris.
BioLocy. Arlé (1933) described nests of 2-12 cells built on rock-faces, just south of Rio de
Janeiro. The cells were invariably horizontal, with a rough mud covering. Cockroach nymphs
were used as prey. Photocryptus apicalis Schmiedeknecht (Ichneumonidae) was a parasite.
Williams (1928 : 115) found two nests near Tena, Ecuador, each fastened to the underside of a
palm leaf near the ground. The prey were cockroaches.
Richards (1937: 109) found two incomplete nests in Guyana, Essequibo River, Moraballi
Creek (c. 30 km SE. Bartica). One was on a tree branch and had six cells (possibly built by two
females), the other was under the roots of a fallen tree and had four cells. Both nests were about
1-3 m from the ground and had the cells ‘mouths upwards’. (This is the only record of vertical
orientation.) The cell building method was identical to that described by Eberhard (1974) for
cameronii. Cockroaches, of which at least one was adult, were used as prey.
Cooper found nests in Colombia: Mocoa attached to a vertical tree trunk; to the underside of
a fallen tree; and to the underside of a palm leaf. The last nest had only just been begun, but the
other two were rather large, with a smooth surface.
Cocoons were similar to those of violascens (cf. Figs 36-37).
A female wasp, taken at Mocoa on 1.vi.1976 by Cooper, was walking over and around several
old and new nests built on the sides and underside of a boulder on the ground. In one cell in each
of two nests was a mixture of cockroach and cricket prey. These were named by Gurney as
134
follows. First cell: Gryllidae, 2 3, 2 9 Anaxipha sp. nr peruviana (Saussure); Blattellidae, 3 nymphs,
all of different species, 1 Chorisoneura sp., 2 gen. indet. Also 1 adult 2 Anaplecta sp. Second cell:
Gryllidae, 1 nymph Eneopterinae, gen. indet.; Blattellidae, 1 ¢ Calhypnorna sp. Only T. grylloc-
tonus has hitherto been reported as preying on crickets. However, it is not certain that the present
wasp was involved with any of the cells near which it was found. Further observation is needed to
establish whether 7. rufiventris also sometimes takes crickets.
MATERIAL EXAMINED. Dates and collectors’ names omitted.
Podium rufiventris Fabricius, holotype 9, GUYANA: Essequibo (Smidt) (UZM, Copenhagen).
Trigonopsis abdominalis Perty, holotype 9, BRAZIL (ZSBS, Munich). Trigonopsis soror Mocsary,
holotype 9, BRAZIL: Amazonas, SAo Paulo de Olivencga (TM, Budapest). Trigonopsis frivaldskyi
Mocsary, holotype 9, BRAZIL: ‘Massanary’ [‘Sao Paulo’ on label], (TM, Budapest).
PERU: 1 9, Junin, Chanchamayo Valley, 800 m (IML, Tucuman); | 9, Junin, Satipo, 750 m
(NM, Rotterdam); 2 9, Loreto, Col. Perené, El Campamento (CU, Ithaca); 1 9, Hudnuco,
Cayumba, 35 km S. Tingo Maria, 800 m (BMNH); 1 9, Loreto, River Pachitea (TM, Budapest);
1 2, Loreto, Atalaya, Upper River Ucayali (BMNH); 10 9, 2 3, Loreto, Pucallpa (BMNH);
1 2, Loreto, Pucallpa, 180 m (Menke coll.); 1 2, Loreto, Pucallpa, 200 m (CMNH, Los Angeles);
1 9, Loreto, Boquer6n Abad (BMNH); | 9, Loreto, Chambireyacu, near Yurimaguas, River
Huallaga (MNHN, Paris); 2 9, Loreto, Rio Tapiche (AMNH, New York); | 3, Loreto, Iquitos
(MP, Belém); 2 9, Loreto, Iquitos, San Roque (CU, Ithaca); 4 9, Loreto, Pebas (3 TM, Budapest;
1 Menke coll.); 1 9, no further data (MNHU, Berlin). EcuApor: 1 6, Napo, Tena (BPBM,
Honolulu); 1 9, Napo, Tena (MNHN, Paris). CoLomsiA: 11 @ (incl. 1 reared), 3 3 (incl. 1 reared),
Putumayo, Mocoa (BMNH); 4 9, 2 3, Putumayo, Alto Afan near Mocoa (BMNH); | 9, Caqueta,
Florencia, 480 m; 4 9, 1 ¢, Meta, Cord. Macarena; 1 2, Amazonas, Leticia; 1 9, Amazonas,
Tarapaca; 3 2, Amazonas, La Chorrera (BMNH); | 9, ‘Bogota’* (MNHN, Paris). VENEZUELA:
1 9, Bolivar, Surukum (CU, Ithaca); 1 9, Bolivar, Canaracuni, 450 m (IZA, Maracay); 1 9, 1 3,
Monagas, Caripito (MCZ, Harvard). GUYANA: 7 9, 1 bred 3, Essequibo River, Moraballi Creek
(c. 30 km SE. Bartica); 2 9, Mazaruni (c. 10 km NW. Bartica), 2nd growth (low forest) (BMNH);
1 2, Bartica District (AMNH, New York); | 9, 1 3, Kartabo, Bartica District (AMNH, New
York); 1 2, same data (MCZ, Harvard); 1 9, Tumatumari (BPBM, Honolulu); 1 9, no further
data (BMNH); | 2, Kamakusa (AMNH, New York). TRINIDAD: | 9, Morne Bleu, 2700 ft (CNC,
Ottawa). SURINAM: 2 9, Coppename River, Raleigh Falls; 1 2, Nassau Mts; 1 9, Lelydorp,
Sumatraweg, Malaise Trap; 1 9, Mapane area; 1 9, Paramaribo (RNH, Leiden); 1 2, Paramaribo
(BMNH); 3 9, no further data (1 RNH, Leiden; 2 MNHU, Berlin). FRENCH GUIANA: 2 2, Mana
River (CM, Pittsburgh); 1 9, ‘Nouveau Chantier’; 1 2, St Laurent du Maroni; | 2, Charvein; 1 9,
“‘Haut-Carsevenne’; | 9, St Georges; 1 9, St Laurent du Maroni (MCZ, Harvard); 6 9, no further
data (2 MNHN, Paris; 2 NM, Vienna; 1 UZM, Copenhagen & 1 FSAE, Gembloux). BRAZIL: 1 g,
Sao Paulo de Olivenca (BMNH); | 2, Amazonas, Tefé (ZSBS, Munich); | 92, 1 3, Amazonas, Villa
Nova (BMNH); | 9, Amazonas, Manaos (MCZ, Harvard); 1 2, 1 $, Amazonas (ZSBS, Munich);
1 9, Amazonas, Rio Purus (NR, Stockholm); 1 3, Amazonas, Fonte Boa (Menke coll.); 1 9,
Amapa, Serra do Navio (Fritz coll.); 22 2, Parad, Belém (4 CU, Ithaca; 6 MP, Belém; 3 ZSBS,
Munich; 2 NM, Vienna; 1 MNHU, Berlin; 1 MIZSU, Turin; 1 MNHN, Paris; 3 BMNH &
1 UM, Oxford); 1 3, Parad, Belém (MP, Belém); 1 9, Para, Belém, Faz. Velha (MP, Belém); 1 9,
Para, Faro (MP, Belém); 1 9, Para, ‘Jabaty’ (BPBM, Honolulu); 5 9, Para, Santarém (2 Menke
coll.; 3 CM, Pittsburgh); 1 9, Para, Maruru near Santarém (CM, Pittsburgh); 1 3, same data
(Menke coll.); 3 2, Amazon, no further data (2 UM, Oxford; 1 NM, Vienna); 1 9, 2 3, Maranhao,
Alcantara (MP, Belém); 2 2, Piaui (TM, Budapest); 1 9, Pernambuco, Recife (MNHN, Paris);
1 2, Bahia, Salvador (NM, Vienna); 1 2, Rio de Janeiro, Niteroi (NM, Rotterdam); | 9, near Rio
de Janeiro (NM, Vienna); 1 2, ‘Monat’ (USNM, Washington); 2 2, no further data (UM, Oxford
& TM, Budapest); 1 3, no further data (MHN, Geneva); 4 9, no data (3 NM, Vienna & 1 UM,
Oxford).
* ‘Bogota’ must here be regarded as an over-generalized locality. The height of this city is well above the upper
altitude limit of the genus.
135
Trigonopsis cameronii (Kohl)
(Figs 1, 40, 69, 70, 98)
[Trigonopsis violaceus Smith; Cameron, 1888 : 26, pl. 2, fig. 13. Misidentification.]
Podium (Trigonopsis) abdominale var. cameronii Kohl, 1902 : 29, 37. Lectotype 2, PANAMA (NM, Vienna),
by designation of Menke in Bohart & Menke, 1976 : 98 [examined].
Trigonopsis cameronii (Kohl); Richards, 1937 : 107, 112, 9.
Lectotype. Menke in Bohart & Menke (1976 : 98) designated as lectotype of cameronii a female
specimen from Panama bearing Cameron’s label ‘Trigonopsis violaceus Smith ?? and Kohl’s
label ‘Tr. abdominalis var. Cameronii’. It is not known how this specimen came to be in NM,
Vienna, but it is part of the original Biologia Centrali-Americana material. This is not discussed
by Menke, and neither are the three paralectotypes (which I have so labelled): 1, a female labelled
‘Bogota’, also bearing Kohl’s labels ‘Trigonopsis abdominalis var. Cameronii K. det. Kohl’ and
‘cf. Trigonopsis violaceus Smith !’; 2, a female from French Guiana bearing Kohl’s label ‘Tri-
gonopsis abdominalis var. Cameronii K. det. Kohl’ and Menke’s labels ‘Syntype: Podium ab-
dominale cameronii Kohl (but not cameronii). Desig. A. S. Menke’ and ‘Trigonopsis violascens
D. T.’; 3, a female from Guatemala which together with the lectotype formed Cameron’s original
Biologia material. It was labelled by him ‘Trigonopsis violaceus’ and is the figured specimen, as
evidenced by the scale line.
2. BL 14-18 (but see below). Almost entirely dark, with violet-blue reflections, especially on head and
gaster. Pronotum without dorsal projection. (Fig. 1.) HW 2:6-3-1, HL 2:7-3-3, HR 0-9-1-0, EL 1-9-2:2,
UIW 1-3-1-5, MIW 1-5-1-7, LIW 1-2-1-4, CTW 0:65-0:80, TOD 0:45-0:60, CR 1-3-1-6, MBL 1-2-1:7,
MAL 0:65-0:95, MR 1-5-2:1, PDL 1-6-2-1, PL 2-4-3-2, PR 1-4-1°6. A very large specimen from Colombia,
Anchicaya differs as follows: BL 21, HW 3-7, HL 3:7, EL 2:5, UIW 1-8, MIW 2:0, LIW 1-7, MBL 1-9,
MAL 1-05, MR 2-4, PL 3-5. Otherwise like rufiventris 9°.
3. BL 14-0-16-0. Colour as 9. Genitalia as in Figs 69, 70. HW 2-6+2°8, HL 2:6-3-0, HR 0-9-1-0, EL 1-8-
2:0, UIW 1-4, MIW 1-5-1-6, LIW 1-2-1-3, CTW 0:25-0:30, CED 0:30-0:32, CER 0-8-1-0, TOD 0°57-0:65,
CR 0-4-0-5, PDL 1-7-2-0, PL 3-0-3-4, PR 1-6—-1-8. (Fig. 40.) Otherwise as rufiventris 3.
This species resembles violascens in colour, but is not sympatric with it. Also, it resembles
rufiventris in structure. However, the constant colour differences, the different averages of varia-
tion as noted below, and their allopatric distribution, added to the fact that only small differences
separate other species of the group, all appear to justify treating cameronii as a good species.
VARIATION. Rarely, one of the female clypeal teeth may be slightly bifid. Only one of the eight
males examined has the inner edge of the mandible with a rather weak preapical expansion, the
others entirely lack it (cf. violascens). This species exhibits less variation than rufiventris, its closest
relative, and the ranges of variation only partly overlap. The average size of cameronii is smaller
and the ranges of clypeal ratio and petiole ratio in the females are not as great. In the males, the
ranges of all the ratios except head ratio are smaller. The genitalia are less variable and on average
smaller and more heavily pigmented.
DisTRIBUTION. Mexico southwards to the west coast of Ecuador, and eastwards to the Sierra
Nevada de Santa Marta on the Caribbean coast of Colombia. It is separated from all other species
of the genus by the Andes (map, Fig. 98).
The species inhabits various types of forest which vary considerably in degree of humidity.
Kohl did not see Smith’s type of violaceus (= violascens) (a normal male), which had not been
illustrated. He correctly referred to violascens, only this type-specimen and an aberrant male. He
described an aberrant female as resplendens, while referring a normal female (from French
Guiana) to cameronii. Thus Kohl had confused the two species. Since he was also misled by a true
cameronii mislabelled ‘Bogota’, he was unable correctly to state the distribution of the two species.
BIOLoGy. Rau (1933 : 182) (Panama, Barro Colorado as) records finding a nest of four cells
on a tree trunk, 0-35 m from the ground.
Eberhard (1972 ; 1974) has given a comprehensive account of this species’ biology, including a
primitive kind of sociality, in Colombia, near Cali. The nests consisted of 1-129 cells, often in-
cluding many old, disused ones, attached to rock-faces. After completion, the cells were incom-
136
e --—~/
ic S |
ev
eft
e265
O) a,
(
. “7
eet aaNet
cameronil Be pan
98
%
La /
Fig. 98 Collection localities of T. cameronii.
pletely covered with rough mud. Cockroaches (Riatia fulgida (Saussure), Chorisoneura translucida
(Saussure) and Amazonina n. sp. (det. Gurney)) were used as prey. Parasites were: Photocryptus
sp. (Ichneumonidae), Gilvella sp. (Tachinidae) (possibly on the roaches) and Macrosiagon lineare
(Le Conte) (Rhipiphoridae).
Cooper found a nest at Colombia, Magdalena, Sierra Nevada de Santa Marta, Rio Don
Diego: of two complete cells, measuring about 25 mm high and 25 mm wide, attached to hanging
rootlets. A wasp was adding pellets to the outside of the nest, giving it a rough appearance.
Cooper informs me that the cocoon of this species resembles that of violascens (Figs 36, 37).
MATERIAL EXAMINED. Dates and collectors’ names omitted.
Podium (Trigonopsis) abdominale var. cameronii Kohl, lectotype 2, PANAMA: Chiriqui, David
(NM, Vienna).
Mexico: | 2, Tabasco, Teapa (BMNH); 1 9, Chiapas, Finca Cucalhuitz, 19 km E. Bochil
(Menke coll.); 2 9, Chiapas, Ruta 190, 21 km E. Cintalapa; 1 2, no further data (USNM, Washing-
ton). GUATEMALA: | 9, Alta Vera Paz, San Juan (paralectotype of cameronii) (BMNH); | 8,
Escuintla, El Salto (BPBM, Honolulu); | 2 no further locality, 300 m (AMNH, New York). EL
SALVADOR: | 9, Los Chorres (CU, Davis). NICARAGUA: | 9, Managua (Menke coll.). Costa RICA:
1 2, Golfito (CMNH, Los Angeles); 2 2, no further data (TM, Budapest). PANAMA: 6 9, Taboga
Island (3 USNM, Washington; 1 TM, Budapest & 2 UZM, Copenhagen); | 3, same data
137
(USNM, Washington); 6 2, Barro Colorado Island (2 CU, Davis; 1 AMNH, New York; 2
USNM, Washington & 1 Menke coll.); 1 9, Chiriqui (TM, Budapest); 1 9, Cerro Campana
(USNM, Washington). CoLomsiA: 3 9, 4 3, Valle, Cali, 1000 m (USNM, Washington); 1 2, same
data (TM, Budapest); 1 2, 1 g, Valle, Rio Jamundi between Cali and Jamundi, 1000 m (BMNH);
1 9, Valle, Cent. Anchicaya, 400 m (BMNH); 2 9, same data (USNM, Washington); 8 9, Narifio,
Barbacoas, 80 m; 2 9, Magdalena, N. Sierra Nevada de Santa Marta, Rio Don Diego (BMNH);
1 2 [Magdalena ?] Minca (CM, Pittsburgh); 1 2, ‘Bogota’ (see footnote p. 135) (paralectotype of
cameronii) (MNHU, Berlin); 1 9, ‘Cananche’ (BMNH); 1 3, ‘Sierra San Lorenzo’ (TM, Budapest).
Ecuapor: | 3, Guayas, Bucay (c. 200 m) (BPBM, Honolulu); 1 2, Balzapamba (1.47 S., 79.13 W.,
c. 200 m) (MNHU, Berlin). 1 9, no data [mislabelled ‘Surinam’] (UZM, Copenhagen).
Trigonopsis howesi sp. n.
(Figs 73, 74, 99)
[ Trigonopsis abdominalis Perty; Howes, 1969 : 36-37, pl. 13b. (Det. Rohwer.) Misidentification.]
2. BL 19-20. Pronotal dorsum without projection, but with a shallow but distinct longitudinal groove.
Gaster red. Petiole of even width. HW 3-4, HL 3-5, HR 1:0, EL 2:4, UIW 1-6, MIW 1-8, LIW 1:6, CTW
violascens
menkei A
howesi x
grylloctonus
Fig. 99 Collection localities of some T. rufiventris-group species.
138
0:85, TOD 0-80, CR 1-1, MBL 1:8, MAL 1:0, MR 1-8, PDL 2:2, PL 3-3-3-4, PR 1-5. Otherwise as rufi-
ventris 2.
3. BL 16:5. Pronotal dorsum with moderately distinct longitudinal groove. Genitalia as in Figs 73, 74.
HW 3-0, HL 3:2, HR 0:9, EL 2:2, UIW 1-5, MIW 1-6, LIW 1:4, CTW 0-32, CED 0-3, CER 1-1, TOD 0°72,
CR 0:4, PDL 2:1, PL 3-4, PR 1-6. Otherwise as rufiventris 3.
This species differs structurally from rufiventris only in its pronotal groove and genitalia.
VARIATION. None observed.
DISTRIBUTION. Guyana only (map, Fig. 99).
BioLoGy. This species builds an extraordinary nest. Howes (1969 : 36-37) states that it was
attached to the underside of a palm frond and consisted of 4-5 horizontal cells placed one beside
each other. These were enclosed in a thin shell-like globe of clay, surmounted by several half-
globes, concave side down, decreasing in size upwards and in contact at least behind. Howes
(pl. 13c) gives a good photograph of one such nest (with four half-globes) and told me in corre-
spondence that he found a total of five such nests. The reason for the extremely elaborate addi-
tions is obscure. Howes states that cockroaches were used as prey.
MATERIAL EXAMINED. Holotype 2, GUYANA: Kartabo near Bartica, 26.ii.1922, reared (Howes)
(abdominalis det. Rohwer) (BMNH).
Paratypes. GUYANA: | 3, same data as holotype but 27.11.1922 (BMNH); 1 9, Mazaruni River,
Kalacoon, 3 mi. from Bartica, 1961 (Howes) (abdominalis det. Rohwer) (USNM, Washington).
Trigonopsis grylloctonus Richards
(Figs 49-51, 75, 76, 99)
Trigonopsis grylloctonus Richards, 1937 : 110. Holotype 3, GUYANA (BMNH) [examined].
Trigonopsis grylloctonus Richards; Menke in Bohart & Menke, 1976: 98.
2. Unknown.
3. BL 13-0. Pronotal dorsum without projection and without longitudinal groove. Gaster red. Petiole
considerably thickened in middle (Figs 49, 50). Genitalia as in Figs 75, 76. HW 2:3, HL 2:3, HR 1-0,
EL 1:6, UIW 1-2, MIW 1-3, LIW 1-0, CTW 0:22, CED 0:20, CER 1-1, TOD 0-50, CR 0-4, PDL 1-4,
PL 2:3, PR 1-6, Fig. 51. Otherwise as rufiventris 3.
The male of this species differs from that of rufiventris only in petiole shape and genitalia.
VARIATION. Unknown.
DISTRIBUTION. Guyana only (map, Fig. 99).
BIoLoGy. Richards (1937 : 111) describes a mud nest found under a leaf, attached to the midrib
near the base. The nest was ovoid, 30 x 20 x 17 mm deep, including the overall mud covering. One
cell was still open and contained a cricket. Later, the male wasp emerged from the only other cell.
This is the only species so far known to select prey other than cockroaches.
MATERIAL EXAMINED. Holotype 3, GUYANA: Essequibo River, Moraballi Creek (c. 30 km SE.
Bartica), 25.ix.1929, reared (BMNH).
The vicina-group
Pronotum with dorsal projection. Propodeum with dorsal rugae also outside the central furrow. Males
with strongly flattened antennae, with some segments expanded. The male clypeal structure of the three
species represents various degrees of development of a basic pattern. Cuspis of genitalia with basal post
round-ended. Female mandible with U- to V-shaped notch, upper edge of apex inflexed. Female clypeus
with 5 usually sharp teeth (centre one occasionally reduced or absent). Sternite 6 sharply keeled.
The females of vicina and schunkei are very similar, having an unusually long third antennal
segment, almost identical clypeal teeth and long, slender mandibles. The female of richardsi is
very different, but its male strongly links it to the other species of the group; in addition, the
female has a pronotal projection; mesoscutal pubescence; and HR of 1-0 in common with all
males of the group.
139
Trigonopsis richardsi sp. n.
(Figs 29-32, 55, 57, 81, 82, 100)
[Trigonopsis affinis Smith; Richards, 1937 : 107. 9 variety, 3. Misidentification. ]
2. BL 13-17. Mandible usually sharply bent at the notch, inner margin + strongly expanded before it,
slender but rather abruptly tapered after it. Upper margin of apex strongly inflexed, especially near notch.
Notch V-shaped, narrow but deep, right-angled distally, shallowly rounded proximally. Clypeus with 2
large teeth and 3 smaller ones between, usually all straight, centre one sometimes reduced or absent.
HW 2:8-3°5, usually exactly equal to HL (at most 0-1 wider in some larger specimens). Occipital carina +
expanded into a thin, lamellar flange, but not as broadly as in rufiventris-group species. (Figs 29-32.)
Pubescence usually dense on anterior half of mesoscutum. HL 2:8-3-4, HR always 1:0, EL 1:8-2:2,
UIW 1:5-1:8, MIW 1-7-2:0, LIW 1-3-1-8, CTW 0-7-1-1, TOD 0-4-0-6, CR 1:2-2:3, MBL 1-2-2:0, MAL
0:3-0:7, MR 2:4-3-4 (— 3-8), PDL 1-7-2:1, PL 2:1-2:7, PR 1-1-1-4.
3. BL 13-15. Antennal segments 2-6 dorsoventrally flattened, 5 & 6 slightly expanded on outer side.
Segment 2 is 1-6-1-7 times longer than wide (Fig. 57). Mandibular inner margin with small but deep basal
emargination. Clypeus with 2 large, sharp teeth, its tooth-to-orbit margin interrupted by a deep notch.
Sides of notch produced into tubercles behind, outer one connected to a third tubercle still further back
(Fig. 55). Genitalia as in Figs 81, 82. HW 2-7-2:8, HL 2:7-2:8, HR always 1-0, EL 1-7-1-9, UIW 1-4-1°5,
MIW 1-6-1:7, LIW 1-3-1:5, CTW 0:40, CED 0:32-0:35, CER 1-1-1-2, TOD 0-65-0-70, CR 0:6, PDL
1:5-1-6, PL 2:3-2:4, PR 1:4-1-5. Otherwise as female, except mesoscutal pubescence not very dense.
The smallest red and black species with a pronotal dorsal projection. Both sexes have a rather
broad occipital flange. Usually the females are well distinguished by mandibular shape, typically
like that of most rufiventris-group species, but variants with weak mandibular development are
more difficult to identify. However, the HR of 1-0 is still distinctive. The male resembles that of
vicina but has fewer antennal segments expanded and three subclypeal tubercles instead of two.
VARIATION. Female variation is complicated and often extreme. The mandibular ratio varies
greatly, independently of specimen size. The mandibular expansion varies from slight to very
strong with a rounded, obtuse angle. When the expansion is slight, the notch is very small. Most
frequently in small specimens, reduction of the centre tooth of the clypeus is accompanied by a
narrowing of the gap between the adjacent pair (cf. Figs 29 & 32). Only in some small specimens,
the tooth-bearing area is very prominent but strongly emarginate, so that the three centre teeth
are effectively much shorter than the outermost pair (Fig. 29). The deeper the clypeal emargina-
tion, the broader and more angular is the mandibular expansion. A small female from Venezuela
entirely lacks the centre tooth and has the tooth-bearing area prominent, but this is not emarginate
and the mandibular expansion is slight (Fig. 30). This specimen is the only one seen which has
the clypeal teeth slightly bent forward. A female from Peru, Pucallpa, 22.vi.1960, has the centre
tooth very reduced and the other four longer than usual, the head strongly expanded posteriorly,
and the mandibular expansion slight (Fig. 31). In this specimen all the interocular widths are
abnormally large for the head width: upper 2-0, middle 1-8, lower 2:0, head width 3-2. Even two
other specimens with head width 3-3 have all the interocular widths smaller (one has the inner
orbits parallel, the other has them slightly divergent downwards). (See also vicina variation.)
If there were not so much material available of this species (nineteen females) it would be easy
to believe that the striking extremes of variation represented distinct species.
DISTRIBUTION. This appears to be mainly a western and northern species, less frequent in the east
(map, Fig. 100). |
BIoLoGy. Unknown.
MATERIAL EXAMINED. Holotype 9, PERU: Loreto, Pucallpa, 28.vi.1960 (Schunke) (BMNH).
Paratypes. PERU: 5 2, same data as holotype but 2, 22.vi.1960; 1, 11.1.1961; 1, 24.xi.1962
(BMNH); 1, 180 m, 19.vi.1971 (USNM, Washington); 2 3, same data but 5.vii.1951 & 6.1.1962
(BMNH); 1 9, Junin, Satipo, 750 m, 15.i.1938 (Lindemans) (NM, Rotterdam); 1 9, Hudanuco,
Tingo Maria, Rio Huallaga, 700 m, 15.i.1947 (Weyrauch) (IML, Tucuman); 1 9, San Martin,
Moyobamba region, 8.i.1926 (Bassler) (AMNH, New York). CoLomsiA: | 2, Vaupes, Miraflores,
400 m, 31.i-5.ii.1972 (Cooper) (BMNH); 2 9, Putumayo, Mocoa, 18.iv.1974 & 31.iii.1976 (Cooper)
140
(BMNH). VENEZUELA: 1 9, Bolivar, Surukum, xii.1940 (Anduze) (CU, Ithaca). GUYANA: 1 9,
Essequibo River, Moraballi Creek (c. 30 km SE. Bartica), 1929 (Richards) (BMNH) (Richards’
affinis variety); 1 9, Bartica (NM, Vienna). BRAZIL: | 9, Piaui (TM, Budapest); 1 3, Amazonas,
Sao Paulo de Olivencga (Bates) (BMNH) (Richards’ ¢ affinis); 1 9, Amazonas, Rio Purus, January
(Roman) (NR, Stockholm). BoLiviA: 2 9, Beni (‘Pando’ on label), Guayaramerin, xii.1956
(Fritz) (Fritz coll.).
Trigonopsis vicina (Dalla Torre)
(Figs 21, 58, 60, 83, 84, 100)
Trigonopsis affinis Smith, 1851 : 31 (part). LECTOTYPE ?, BRAzit (BMNH), here designated [examined].
[Junior secondary homonym in Sceliphron Klug of Sphex affinis F., 1793.]
Sceliphron vicinum Dalla Torre, 1897 : 392. [Replacement name for Trigonopsis affinis Smith.]
Trigonopsis affinis Smith; Richards, 1937 : 107, 111, 2 (part).
LECTOTYPE DESIGNATION. Smith described affinis from specimens collected by Wallace and Bates.
One syntype in the BMNH collections agrees with Smith’s description and locality, and with the
current interpretation of the species. A second syntype in the collections of UM, Oxford, does not
agree with the description or locality, and is identified as the species here described as cooperi. I
have labelled, and here designate as lectotype, the female in the collections of the BMNH.
?. BL 16-19. Mandible slender, evenly curved. Mandibular notch formed by an oblique incision where
the inflexed upper margin of the apex is twisted under the upper margin of the basal part, with the basal
angle of the notch more shallowly obtuse than the distal. Clypeus with 5 usually well-formed teeth,
outermost pair the longest, the centre one equal to or slightly longer than the adjacent pair (Fig. 21).
Antennal segment 3 is 2:5-2:8 times as long as 6. MIW 2:2-2-4 times CTW. HW 3-1-3-7, HL 2:9-3-4,
HR always 1:1, EL 1-9-2:2, UIW 1-6-1:9, MIW 1-8-2:2, LIW 1-5-1:8, CTW 0-8-0-:9, TOD 0-5-0-7,
CR 1-3-1-6, MBL 1-7-2-1, MAL 0-3-0-7, MR 2:7-3:2 (— 5:6), PDL 1-7-1-9, PL 2:5-2°8, PR 1:4-1°6.
$. BL 16:5-19-0. Antennal segments 2-12 rather strongly dorsoventrally flattened, segments 5-10
strongly expanded on outer side. Segment 2 about as long as wide (Fig. 58). Extreme base of mandibular
inner margin scarcely emarginate. Clypeus with 2 large, sharp teeth. Base of outer side of each tooth
tuberculate behind. Behind the tooth-to-orbit margin, but not forming part of it, is another tubercle
whose tip is just visible in front view. Together with the other tubercle it forms the sides of a deep notch.
There is no third tubercle behind the outer one (Fig. 60). Very similar to male neotropica (Fig. 54).
Occipital fossa width 1-2 (-1-8). Eyes convergent downwards. Genitalia as in Figs 83, 84. HW 3-0 (4-1),
HL 3-0 (-3-8), HR 1-0(-1-1), EL 1:9 (—2:4), UIW 1-6 (-2:2), MIW 1:8 (-2-4), LIW 1-4 (2:2), CTW
0-35 (-0-80), CED 0-35 (-0-60), CER 1-0 (-1:3), TOD 0-75 (0:90), CR 0:5 (-0:9), PDL 1-8 (-2:3),
PL 2:9 (-3-2), PR (1:4-) 1-6. Otherwise as female.
The female is similar to that of schunkei, the most obvious difference being the much greater
MR of vicina. The male is very like that of neotropica in clypeal structure, but differs mainly in
that some of the antennal segments have lateral expansions.
VARIATION. A large female from Trinidad, although its head width (3-5) is not the largest, has the
head posteriorly expanded more than that of any other specimen, and is reminiscent of the
Pucallpa specimen of richardsi (p. 140). Apart from this and the central clypeal tooth, there is
little variation in females.
In the absence of more specimens better to show the range of variation, the male from Marajé
is placed here only tentatively. Although its genitalic structure readily distinguishes it from
neotropica, the structure of the clypeus and antennae of this very large individual are modified
such as to make its separation from richardsi difficult.
DISTRIBUTION. Sparingly from the Amazon northwards (map, Fig. 100).
BioLoGy. Unknown. Williams reared a male in 1923 (the only reared specimen of the species-
group) but regrettably gave no details in his 1928 paper.
MATERIAL EXAMINED. Trigonopsis affinis Smith, lectotype 9, BRAZIL: Parad, Belém (Wallace &
Bates) (BMNH).
141
richardsi
vicina
schunkei
100
Fig. 100 Collection localities of the 7. vicina-group.
COLOMBIA: 1 9, Vaupes, Mitt, 17.v.1974 (Cooper) (BMNH). Ecuapor: 1 9, Napo, Napo,
viii-ix.1931 (Benoist) (MNHN, Paris). TRINIDAD: 1 9, Diego Martin, 21.i.1942 (McC. Callan)
(USNM, Washington); 1 9, Arima Valley, 21.vi.1951 (Menke coll.). GUYANA: 1 9, ix.1923; 1 3,
reared 12—13.x1.1923, Blairmont Plantation, Berbice County (Williams) (BPBM, Honolulu); 1 9,
Pakaraima Mts, Upper Ireng River, 1932 (Myers) (BMNH); 1 9, Demerara (MCZ, Harvard).
SURINAM: | 9, Paramaribo, Charlesburg, 16-18.i.1964, Malaise trap (Geijskes) (RNH, Leiden).
BRAZIL: 1 3, Para, Ilha do Marajé, R. Anajas, 13.vi.1900 (Ducke) (MP, Belém).
Trigonopsis schunkei sp. n.
(Figs 26, 52, 56, 79, 80, 100)
2. BL 18-22. Mandible slender and evenly curved. Mandibular notch only slightly oblique, inflexed upper
margin of apex cut off perpendicularly well short of rounded end of upper margin of base, so that a broad,
deep notch is formed. Antennal segment 3 is 2-:9-3-3 times as long as 6. MIW 1:-7-2:1 times CTW.
Clypeus with 5 usually well-formed teeth, the outermost pair the longest and the centre one equal to or
shorter than the adjacent pair (Fig. 26). HW 3-7-4-4, HL 3-1-3-6, HR 1-1-1-2, EL 2:0-2-4, UIW 1-9-2:2,
MIW 2:-1-2°5, LIW 1-9-2:3, CTW 1-1-1:2, TOD 0-6-0-:7, CR 1-6-2:1, MBL 2-1-2:°6, MAL 1-0-1-2,
MR 1:9-2-:3, PDL 2:0-2-4, PL 3-0-3-3, PR 1-3-1°5.
142
3. BL 15-:0-16°5. Antennal segments 2-8 strongly flattened dorsoventrally, segments 4-8 moderately
expanded on outer side. Segment 2 a little longer than wide (Fig. 56). Inner mandibular margin with a
sub-basal tooth. Clypeus produced into two large, rectangular, divergent lobes, with a shallow emargina-
tion at the end of each, and a broad, deep one between. Lobe-to-orbit margin with a deep notch, the lobe
side of which is strongly carinate behind, with an unconnected tubercle further back (only visible in ventral
view) (Fig. 52). Genitalia (Figs 79, 80). HW 3-1-3-5, HL 3-1-3-5, HR 1-0, EL 1-9-2-1, UIW 1-8-2-0,
MIW 1-9-2-:2, LIW 1-7-2:0, PDL 1:7-1-8, PL 2:3-2-4, PR 1-3. Otherwise as female except pubescence
very dense on anterior half of mesoscutum.
The female is very similar to that of vicina but has a much smaller mandibular ratio as well as
other more detailed differences. The male has a unique clypeal shape.
VARIATION. Apart from the centre tooth of the female clypeus, the back of the head in the Pucallpa
1.v.1959 male is very strongly expanded, that of the other one scarcely so.
DISTRIBUTION. Amazon Basin and Guianas; uncommon (map, Fig. 100).
BioLocy. Unknown. Cooper noted an offensive smell produced by the female of this species when
caught, although it was less strong than that of succinea.
MATERIAL EXAMINED. Holotype 92, PERU: Loreto, Pucallpa, 3.vii.1960 (Schunke) (BMNH).
Paratypes. PERU: 1 9, same data as holotype but 20.ii.1963 (BMNH); 2 3, same data as holotype
but 1.v.1959 (BMNH) and 2.iii.1948 (USNM, Washington); 1 9, Loreto, Iquitos, San Roque,
iv.1929 (Klug) (CU, Ithaca). CoLomBiaA: | 9, Meta, La Macarena 29.x-7.xi.1976 (Cooper)
(BMNH). Guyana: | 2, Mazaruni (c. 10 km NW. Bartica), 2nd growth (low forest), 27.xi.1937,
no. 346 (Richards & Smart) (BMNH). BRraziL: 2 9, Amazonas, Tefé (Bates) (ZSBS, Munich); 1 9,
Para, Belém, 1846 (Ghiliani) (MIZSU, Turin); 2 9, Para, Belém, 6.v.1902 & 23.vi.1903 (Ducke)
(MP, Belém).
The intermedia-group
Pronotum with sharply angulate dorsal projection. Propodeum with dorsal rugae also outside central
furrow. Female mandible very stout, nearly straight and parallel-sided almost to the U-shaped notch.
Notch always large, mandibular outer margin strongly curved around it. Upper margin of apex broadly
inflexed, especially near the notch, abruptly tapered. Clypeus with 2 large teeth and 4 or 5 smaller, blunt,
often irregular teeth between. Male clypeus with 2 large, sharp, simple teeth, and the tooth-to-orbit
margin entire. Antennal segments cylindrical. Cuspis of male genitalia with basal post round-ended.
The females of intermedia and cyclocephalus are very similar. They are larger than moraballi,
with the clypeal teeth more numerous and more irregular, the mandible less strongly curved, its
apex less slender, the notch overhung by a carina and the upper edge of both base and apex
inflexed.
Trigonopsis intermedia Saussure
(Figs 14, 15, 62, 63, 87, 88, 101)
Trigonopsis intermedius Saussure 1867 : 33, pl. 2, fig. 18. Holotype 2, BRAziL (NM, Vienna) [examined].
[Podium (Trigonopsis) affine Smith; Kohl, 1902 : 34, part. Misidentification.]
9. BL 22-27. The largest known species of the genus, and the only species lacking a basal spine on the
mandibular inner margin (reduced to an inconspicuous tubercle). Mandible very stout, most of the base
slightly curved and parallel-sided, strongly curved from shortly before the notch to beyond it. Near the
notch, apical upper margin broadly inflexed, basal very narrowly. Notch large, right-angled distally,
obtuse proximally, partly overhung by a strong carina. Clypeus with the outermost pair of teeth fairly
large, outwardly directed and strongly bent forward. Between them are 4 much smaller, often irregular,
teeth. The central gap (usually the widest) sometimes has a vestigial, blunt, seventh tooth (Figs 14, 15).
HW 4-4-5:2. HL 3-44-0, HR 1-2-1-3, EL 2-5-3-0, UIW 2-0-2:3, MIW 2-5-2:9, LIW 2:3-2:7, CTW
1-5-1:8, TOD 0-5-0:7, CR 2-6-3-4, MBL 2-4-3-3, MAL 0°7-1:1, MR 2-6-3-4 (-3-9), PDL 2°5-3-1, PL
2°6-3-2, PR 1-0-1:1. ;
3. BL 19-5-23-0. Mandibular inner margin with sub-basal tooth. Clypeus with the 2 teeth widely sep-
arated, their tips often bent outwards. Head + strongly expanded posteriorly (Figs 62, 63). Genitalia as
143
in Figs 87, 88. HW 3-9-4-5, HL 3-5-4-1, HR 1-1, EL 2:4-2:7, UIW 1-9-2:3, MIW 2:-3-2:7, LIW 2:2-2°5.
CTW 0:90-1:05, CED 0:52-0:67, CER 1:4-1:7, TOD 0:92-1:10, CR 0-9-1-1, PDL 2:2-2°8, PL 2-9-3-6,
PR 1-1-1-4. Otherwise as female.
The largest species. The female is very similar to that of cyclocephalus, but lacks the mandibular
spine. The males of both this species and schunkei have a mandibular tooth, but the clypeal shape
is quite different.
VARIATION. The temples of the male may be + strongly expanded, and the tips of the clypeal
teeth are sometimes bent outwards. The clypeal teeth of females are irregular as detailed above.
DISTRIBUTION. Fairly common in the Guianas, less so on the eastern coast of Brazil (as far as
Rio de Janeiro ?) (map, Fig. 101).
TYPE-LOCALITY. The original description and label both give only Brazil. According to the report
of the voyage, the ship Novara called in Brazil only at Fernando Noronha Island and Rio de
Janeiro. The latter seems the most likely to be the type-locality, especially since at that time the
coastal rain-forest was continuous with that further north, where all other specimens were
collected.
BIoLoGy. Richards (1937 : 107) states that the female taken at Moraballi Creek on 22.x.1929 was
carrying a cockroach up a hollow in a Mora tree.
The following notes were made in Guyana by Professor Richards during the 1937 expedition.
They refer to nests examined or collected at Mazaruni (c. 10 km NW. Bartica). ‘2 nests 19. viii. 1937,
the first on underside of sloping, living tree, 1 ft from ground, in secondary growth. 2 cells,
1 empty, the other with a wasp. The second nest about an inch from the first, 4 cells.’
‘Several groups of cells 25.viii.1937 on buttress of old Mora tree. First group very old, occupied
by termites. Second group, 2 cells, | with a cockroach, with an egg in front of mid coxae, the other
with 2 cockroaches and a larva 10 mm long (one prey still intact, constantly moved all its palpi).
Third group, 3 cells, one with 4 cockroaches. Preserved. [Now lost.] One cockroach with a larva
4 mm long between fore and mid coxae. All more or less alive, one almost able to walk. Another
cell with a larger larva, another with a cocoon. Fourth group, 2 cells. Both with cocoons.’
The above nests produced wasps as detailed in ‘Material examined’, both sexes from some
nests.
MATERIAL EXAMINED. Holotype 2, BRAZIL, (Rio de Janeiro ?) (Novara Expedition) (NM, Vienna).
GUYANA: 2 9, Essequibo River, Moraballi Creek (c. 30 km SE. Bartica), 29.viii & 22.x.1929
(Oxford University Expedition) (BMNH); 1 9, 2 3, Mazaruni (c. 10 km NW. Bartica), 2nd growth
(low forest), 19.vili.1937, reared (Richards & Smart); 3 2, 2 3, same data but 25.viii.1937, reared
(BMNH); | 3, Demerara (Ballion) (IRSNB, Brussels); 1 9 (Rodway); 2 2 (Bartlett); 1 $ (Cameron),
all without further data (BMNH). SuRINAM: 2 2 without further data (1, FSAE, Gembloux; 1,
TM, Budapest). FRENCH GUIANA: | 9, 1 3, Mana River, June 1917 (CM, Pittsburgh); 1 9, St
Jean de Maroni; | 9, ‘Nouveau Chantier’; 1 9, Charvein (MNHN, Paris); 1 9, Les Hattes, Bas
Maroni (Le Moult) (Menke coll.); 1 ¢ ‘Laiord’ [?] (MNHU, Berlin); 1 9 (Ballion) (IRSNB,
Brussels); 1 2 without further data (UZM, Copenhagen); 1 3 without further data (IRSNB,
Brussels). BRAZIL: 1 9, Para, Belém, 1846 (Ghiliani) (MIZSU, Turin); 1 9, Para, Belém, 8.vii.1902
(Ducke) (MP, Belém); 1 9, no locality (Ballion) (IRSNB, Brussels); 1 9, 1 3, without data (NM,
Vienna); 2 3 without data (FSAE, Gembloux).
Trigonopsis cyclocephalus Smith
(Figs 22, 23, 38, 39, 64, 66, 89, 90, 101)
Trigonopsis cyclocephalus Smith, 1873 : 54. Holotype 3, BRAzIL (BMNH) [examined].
[Podium (Trigonopsis) affine Smith; Kohl, 1902 : 34, part. Misidentification.]
Trigonopsis cyclocephalus Smith; Menke in Bohart & Menke, 1976: 98.
2. BL 20-22. Mandible very stout, the base slightly curved and parallel-sided almost to the notch, where
it is strongly curved. Near the notch, the upper margin of the base is very narrowly inflexed, that of the
144
apex broadly. Notch large, right-angled to obtuse distally, more obtuse proximally, and partly overhung
by a strong carina. Outermost pair of clypeal teeth large, well-formed, strongly bent forwards, sometimes
notched near the base of the outer side, but the tips not bent back. Between them are usually 4 much
smaller and often irregular teeth with a wide central gap, but occasionally a short but very broad seventh
tooth fills the latter; some teeth may be fused, occasionally asymmetrically. (Figs 22, 23.) Pronotum
with sharply angulate dorsal projection. Gaster red. HW 4:3-4-5, HL 3-3-3-6, HR 1:2-1-3, EL 2:4-2°6,
UIW 1-:9-2-1, MIW 2:4-2:6, LIW 2:2-2:4, CTW 1:4-1:5, TOD 0:6-0-8, CR 1-7-2:5, MBL 2-5-2-8, MAL
0-8-1:1, MR 2-4-3-3, PDL 2:5-2:6, PL 2:4—2:8, PR 1-0-1°1.
3. BL 15-19. Clypeus with the 2 teeth narrowly spaced (Fig. 64). Third antennal segment 1-1—1-3 long,
1-:8—1-9 times as long as the first. Sternite 8 with rather sharp keel near the tip, which is narrowly truncate
(Fig. 66). HW 3-1-3-7, HL 2-8-3-2, HR 1:1-1:2, EL 1-8-2:3, UIW 1-6-1-:8, MIW 1-9-2:2, LIW 1-6-1-9,
CTW 0:35-0:40, CED 0:35-0:40, CER 0:9-1-1, TOD 0:85-0:97, CR 0-4-0:5, PDL 1-8-2:2, PL 2:3-3-0,
PR 1-2-1-4. Otherwise as female.
The female is very similar to that of intermedia, but has a mandibular spine. The male is
extremely similar externally to those of moraballi and cooperi. However, cooperi is easily dis-
tinguished by its genitalia, while the differences between those of moraballi and cyclocephalus are
numerous but slight. See keys (p. 129).
THE NON-TYPE SPECIMENS. The males from Leticia reared by Cooper differ from the holotype
male in that they are smaller, their temples less expanded and their genitalia slightly different.
However, I feel that they are better placed with cyclocephalus than with moraballi. The identi-
fication of female cyclocephalus as here treated is based largely on this reared material.
VARIATION. Back of male head + strongly expanded. Female clypeal teeth irregular as detailed
above.
DISTRIBUTION. So far known only from the Upper Amazon (map, Fig. 101).
BIOLOGY. Two mud nests, each of four cells, were found by Cooper at Leticia on 31.viii.1974,
7°6 cm apart, under a flattened liana. One cell of the first was filled with cockroaches (Blaberidae,
1 2 Poroblatta or gen. near, perhaps n. sp. (det. Gurney)), a second contained a full-grown larva,
and the remainder, pupae. The other nest had an entirely smooth surface, i.e. the cells had a
complete, overall covering. It measured 750 mm long by 350 mm wide and was roughly oval in
shape, but distinctly smaller at one end [perhaps with male cells ?]. Both sexes were reared from
these nests. Cooper found two other nests at La Chorrera, Colombia. The first, on 25.viii.1976,
consisted of a single cell 270 mm by 170 mm, with a rough surface, under a leaf 2:5 m from the
ground. A male wasp was reared. The second nest, on 28.viii.1976, had two cells, measured
40 x 35 x 15 mm, and had a smooth surface. It was under a leaf 2 m from the ground. The first
cell contained a larva which failed to develop, the second was still being provisioned by a female,
which was captured. The cockroaches were | 3, 1 2, of the same species as those from the Leticia
nest (det. Gurney).
Cooper informs me that all the cocoons of cyclocephalus he found resembled those in Figs 38,
39.
MATERIAL EXAMINED. Holotype 3, BRAZIL: Amazonas, Tefé (Bates) (BMNH).
BRAZIL: 1 9, Amazonas, SAo Paulo de Olivenca (UM, Oxford); 1 9, same locality (Bates)
(ZSBS, Munich); 2 2, Amazonas, Tabatinga, 7 & 8-9.vili.1974 (Cooper) (BMNH). CoLosia: | 9,
Amazonas, Leticia, 19-20.viii.1974; 3 9, 2 3, same locality, 31.viii.1974, reared (Cooper); 1 3,
Amazonas, Rio Igara Parana, La Chorrera, 25.vili.1976, reared (Cooper); 1 9, same data but not
reared; 1 9, same data but 28.viii.1976, not reared (BMNH).
Trigonopsis moraballi Richards
(Figs 12, 13, 61, 67, 91, 92, 101)
Trigonopsis moraballi Richards, 1937 : 107, 112. Holotype 2, GUYANA (BMNH) [examined].
?. BL 19-21. Mandible very stout, parallel-sided and straight almost to the notch, where it is very strongly
curved. Apex very slender, upper margin broadly inflexed, especially near the notch, but that of the base
not at all so. Notch large, right-angled or acute distally, rounded proximally, not overhung by a carina.
145
intermedia ®
cyclocephalus m=
moraballi x
Fig. 101 Collection localities of the 7. intermedia-group.
The large, outermost pair of clypeal teeth strongly bent forward, tips slightly bent back, notched on outer
side near tip. Between them are 4 smaller teeth, fairly regular in height and spacing, but centre pair more
obtuse, sometimes slightly shorter and closer together (Fig. 12). HW 4:1-4:2, HL 3-2-3-3, HR 1-2-1°3,
EL 2:3-2:4, UIW 1-9-2:0, MIW 2:3-2:5, LIW 2-0-2:1, CTW 1-2-1:3, TOD 0-5-0-6, CR 2-1-2-4, MBL
2°3-2:5, MAL 0-5-0-8, MR 2:9-3-1 (4-4), PDL 2:-1-2:2, PL 2-4-2:6, PR 1-1-1-2.
3. BL 16. Clypeus with the 2 teeth narrowly spaced (Fig. 61). Third antennal segment 1-0 long, 1-6
times as long as the first, MIW 0-35 more than UIW. Sternite 8 with moderately sharp keel, its tip broadly
rounded (Fig. 67). Genitalia as in Figs 91, 92. HW 3:2, HL 2:8, HR 1:1, EL 2:0, UIW 1-6, MIW 2:0,
LIW 1-5, CTW 0:35, CED 0-32, CER 1:0, TOD 0-70, CR 0-4, PDL 1-7-1-8, PL 2:4-2-5, PR 1-4. Otherwise
as female.
The female is rather like those of intermedia and cyclocephalus, but its mandibular notch lacks
an overhanging carina and the clypeal teeth are less irregular.
The male is extremely similar externally to those of cooperi and cyclocephalus. However,
cooperi has distinct genitalia, while the differences between those of moraballi and cyclocephalus
are numerous but slight (see keys p. 130).
VARIATION. In the single female without locality the 2 centre teeth are much shorter than the
adjacent pair, one of them is displaced outwards and the larger gap thus created has a blunt,
vestigial seventh tooth (Fig. 13). These differences suggest that this specimen may not belong to
146
moraballi, but in the absence of other evidence it seems best to regard it as representing variation
within the species.
DISTRIBUTION. So far known only from Guyana, but the much paler colour of the specimen without
data suggests the possibility of an Upper Amazon origin (map, Fig. 101).
BIoLoGy. Richards (1937 : 108-109) gives a photograph of a 2-celled nest and notes on others. He
states that they were found under leaves, 1-2—1-8 m from the ground. Each of the mud cells had
one end touching the leaf mid-rib. There was a maximum of 3 cells per nest, the cells being parallel
to and in contact with each other. Cockroaches were used as prey, and a wasp was seen to carry
one with its ‘mouth and forelegs’. A bombyliid fly, Anthrax leucopyga Macquatt, was also reared.
MATERIAL EXAMINED. Holotype 9, GUYANA: Essequibo River, Moraballi Creek (c. 30 km SE.
Bartica), 2.ix.1929, reared (Oxford University Expedition) (BMNH).
GUYANA: | 9, same data as holotype; 1 9, same data except 3.x.1929, not reared; 2 J, same
data except 13.vili.1929, not reared, and 20.ix.1929, reared (BMNH) (all paratypes of moraballi).
1 2 without data (UM, Oxford).
The neotropica-group
All species are uniquely characterized by: female mandibular notch formed by a step-like thickening,
and cuspis of male genitalia with apex of basal post truncate. Further characters are: pronotum with
dorsal projection, propodeum with dorsal rugae also outside central furrow, female clypeus with 5 teeth.
Within the group, cooperi is distinguished by: slender female mandible, notch vestigial (never
absent), and cylindrical male antennae. The remaining species are further united by: males with
antennal segments slightly flattened, and offset at their junctions, females with identical mandi-
bular shape unique to this group: rather stout, slightly curved towards tip, inner margin slightly
expanded before notch. The females of two species are still further united by: outermost pair of
clypeal teeth equal to or slightly shorter than adjacent pair, another character unique to this
species-group.
Rugae outside central furrow of propodeal dorsum often markedly obsolescent anteriorly,
especially in succinea and cooperi.
The evidence so far available suggests that the species of this group, unlike the others in the
genus, add no extra mud to the nest after the cells are complete. It is curious that, despite this,
extreme mandibular wear is found in an unexpectedly high proportion of females (see also
succinea biology).
Trigonopsis neotropica sp. n.
(Figs 18-20, 27, 54, 85, 86, 102)
[Trigonopsis cyclocephalus Smith; Richards, 1937 : 107, putative 9. Misidentification.]
2. BL 19-20. Mandible stout, inner margin of base a little expanded before the step-like notch. Small,
basal spine of mandibular inner margin narrow, pointed (Fig. 18). Clypeal teeth + bent forwards, the
outermost pair most strongly, often slightly broader and more blunt than adjacent pair. Centre tooth
smaller than others. Centre pair of gaps a little narrower than outer (Figs 18-20). Occipital carina often +
expanded into a brown, translucent lamellar flange. Pronotal dorsal projection scarcely concave anteriorly
(Fig. 27). Wings not amter-tinted, stigma and most veins dark brown. HW 3:7-3:9, HL 3-2-3-5, HR
1-1-1-2, EL 2-2-2:4, UIW 1-7-1-8, MIW 2:1-2-3, LIW 1-8-2:0, CTW 1-2-1-4, TOD 0-45-0-50, CR 2-7-2°9,
MBL 2:0-2:2, MAL (0-25-) 0:45-0:60, MR 3-3-4-4 (-8-4), PDL 2:2-2:4, PL 2:9-3-2, PR 1:2-1-4.
3. BL 18. Clypeus with 2 large, sharp teeth. Base of outer side of each tooth is tuberculate behind.
Behind tooth-to-orbit margin but not forming part of it, is another tubercle whose tip is just visible in
front view. Together with the other, it forms the sides of a deep notch. There is no third tubercle behind the
outer one (Fig. 54). Clypeus similar to that of male vicina (Fig. 60). Inner orbits parallel. Antennal seg-
ments slightly flattened dorsoventrally, their junctions offset in lateral view (cf. Fig. 59). Occipital fossa
width 1-5. Genitalia as in Figs 85, 86. HW 3-6, HL 3-3, HR 1-1, EL 2:2, UIW 1-8, MIW 2:1, LIW 1:8,
CTW 0:65, CED 0-45, CER 1-4, TOD 0-82, CR 0:8, PDL 2:1, PL 2:9, PR 1-4. Otherwise as female.
147
neotropica
succinea x
mocoana A
cooperi ®
Fig. 102 Collection localities of the 7. neotropica-group.
The female is similar to that of succinea, but is distinguished by its smaller size, the basal spine
of the mandibular inner margin pointed instead of rounded, shorter petiole, less coarse propodeal
sculpture etc. The male is very like that of vicina, but lacks the antennal expansions.
VARIATION. The Belém (i.e. easternmost) specimens differ from the others in the following ways.
The body is altogether darker, in particular the clypeus, legs and wing-bands.
The second abscissa of the radial vein is markedly shorter than the third (roughly equal in the
other specimens).
In the females, the central pair of gaps between the clypeal teeth are shallower than the outer
pair (Fig. 20), whereas they are roughly equal in the other females (Figs 18,19).
Although all these forms of individual variation are usually found separately in various other
species of the genus, it is interesting that they occur together, as well as being geographically
correlated, in neotropica.
DISTRIBUTION. Widespread but uncommon (map, Fig. 102).
BioLoGy. The paratype female bears a label ‘From one of 3 clay cells fastened side by side under
arboreal termites nest. One cell already vacated and one pupa broken in fall.’ Hamilton further
informs me that the nest was about 5m from the ground, and that the individual cells were
visible.
If this is a typical nesting-site for the species-group (excepting the aberrant species cooperi)
148
they may build their nests at a much greater height than other Trigonopsis. This would help to
explain why it is much the rarest species-group in collections (see also succinea biology).
MATERIAL EXAMINED. Holotype 9, BOLIVIA (Steinbach) (BMNH) (Richards’ putative cyclocephalus
9).
Paratypes. PERU: 1 9, Hudnuco, Tingo Maria, 20-27.i.1968 (Garcia & Porter) (MCZ, Harvard).
BRAZIL: 1 9, Mato Grosso, Serra Roncador, Royal Society Base Camp, 12.50S., 51.47 W.,
11.vi.1968, reared (Hamilton) (BMNH); | 9, 1 3, Para, Belém, 6.iii.1900 (Ducke) (NM, Vienna);
1 9, Para, Belém, 30.vi.1900 (Ducke) (MP, Belém).
Trigonopsis succinea sp. n.
(Figs 24, 25, 28, 53, 59, 95, 96, 102)
?. BL 22-26. Mandibular base with inner margin a little expanded before the step-like notch. Small basal
spine of mandibular inner margin rather broad, rounded at tip. Outermost pair of clypeal teeth scarcely
shorter but + broader and blunter than adjacent pair. Centre tooth equal to or shorter than outermost
pair. Teeth + asymmetrical. Centre pair of gaps wider and deeper than outer pair. Lower part of clypeus
and teeth a little bent forward (Figs 24, 25). Occipital carina slightly lamellar flange-like, black, trans-
lucent only when seen against the light. Pronotum with dorsal projection strongly transverse, in profile
very strong and sharply angulate with distinct concavity in front (Fig. 28). Pubescence of anterior half of
mesoscutum + dense. Wings + strongly amber-tinted, stigma pale (much paler than infuscation of radial
cell). Rugae outside central furrow of propodeal dorsum very coarse, rather widely spaced, and usually
strongly obsolescent anteriorly. Central groove very deep throughout. HW 3-8-4-6, HL 3-5-4-0, HR
1-1-1:2, EL 2-3-2-7, UIW 1-8-2:1, MIW 2:2-2-6, LIW 1-9-2:3, CTW 1-1-1-4, TOD 0-6-0-7, CR 1-8-2:2,
MBL 2:1-2:5, MAL 0-45-0-80, MR 2:8-4-4 (-4-9), PDL 2-5-3-1, PL 3-7-4-4, PR 1-3-1°5.
$. BL 17-20. Clypeus with two large, sharp teeth. Tooth-to-orbit margin entire, without tubercles
(Fig. 53). Antennal segments slightly dorsoventrally flattened, their junctions offset in lateral view (Fig.
59). Head + expanded posteriorly. Genitalia Figs 95, 96. HW 3-3-3°8, HL 3-0-3-5, HR 1-1, EL 2-0-2:2,
UIW 1:7-2:1, MIW 1-9-2-2, LIW 1-6-1-8, CTW 0:37-0:60, CED 0:30-0:47, CER 1:2-1-3, TOD 0:77-0:87,
CR 0:5-0-7, PDL 2:0-2:4, PL 2:7-3-8, PR 1-3-1-:6. Otherwise as female, except occipital carina narrower.
The female is similar to that of neotropica, but the basal spine of the mandibular inner margin
is rounded instead of sharp, the insect is larger, has a longer petiole, coarser propodeal sculpture,
etc. At first sight the male much resembles those of several other species, but has slightly flattened
antennae. The identification should always be confirmed by genitalia examination (Figs 95, 96).
VARIATION. Female clypeal teeth as detailed above. The back of the head of the male from Tena
is very strongly expanded, that of the one from Riberalta scarcely so.
DISTRIBUTION. East Andean foothills from Colombia to Bolivia. Most records are from about
600 m, with isolated ones down to about 200 m (map, Fig. 102).
BIOLOGY. Cooper has observed females collecting various kinds of mud (on one occasion in the
company of a rufiventris female). All of them flew high into the forest, which strengthens the
supposition (discussed under neotropica) that most species of this group build their nests far
above the ground.
Cooper informs me that the females especially of this species emit an offensive smell when
captured.
MATERIAL EXAMINED. Holotype 2, EcUADoR: Napo, Tena, 14.iv.1976 (Cooper) (BMNH).
Paratypes. ECUADOR: 2 9, same data as holotype but 16 & 19.iv.1976 (BMNH); | 3, Napo,
near Tena, v.1923 (Williams) (BPBM, Honolulu). CoLomsia: 2 9, Putumayo, Mocoa, 580 m,
31.iii., 2.vi.1976 (Cooper) (BMNH); 1 9, same data but 5.vi.1976 (USNM, Washington). PERU:
1 9, Loreto, Pucallpa, 14.ii.1952 (Schunke) (BMNH). BottviA: 1 9, “Songo’ (TM, Budapest); 1 3,
Beni (‘Pando’ on label), Riberalta, xi,1956 (Fritz) (Fritz coll.).
Trigonopsis mocoana sp. n.
(Figs 35, 102)
9. BL 18-20. Mandible stout, inner margin of base a little expanded before the notch, which is step-like
and very small (but not vestigial as in cooperi). Outermost pair of clypeal teeth the longest. Centre tooth
149
slightly shorter than adjacent pair, and about half height of outermost pair and narrower than them. All
teeth bent forward, outermost pair most strongly. All gaps are equally deep but centre pair slightly
narrower than outer pair (Fig. 35). Pronotal dorsum with sharply angulate projection. Pubescence mod-
erately dense on anterior half of mesoscutum. Gaster red. HW 3-6—-3-8, HL 3-0-3-2, HR 1-2, EL 2-2-2°3,
UIW 1-7, MIW 2:-1-2:2, LIW 1-7-1-8, CTW 0-9-1-0, TOD 0-6-0-7, CR 1-:3-1:7, MBL 1-9-2:0, MAL
0:5-0-6, MR 3-4-3-8, PDL 2:2-2:4, PL 2-5-2:6, PR 1:1.
3. Unknown.
The female is rather like that of cooperi, but is distinguished by the mandible having a larger
notch, with the inner margin expanded before it; and by the smaller clypeal ratio.
VARIATION. Slight irregularity in female clypeal teeth.
DISTRIBUTION. Known only from the one locality (map, Fig. 102).
BioLocy. Unknown.
MATERIAL EXAMINED. Holotype 2, COLOMBIA: Putumayo, Mocoa, 24.xii.1974 (Cooper) (BMNH).
Paratype. 1 9, same data as holotype but 6.iv.1976 (BMNH).
Trigonopsis cooperi sp. n.
(Figs 33, 34, 65, 68, 93, 94, 102)
[Trigonopsis affinis Smith, 1851 : 31, 2, part. Misidentification.]
[Trigonopsis moraballi Richards, 1937 : 107, 3, part. Misidentification. ]
[Trigonopsis cyclocephalus Smith; Richards, 1937 : 107, 3, part. Misidentification.]
?. BL 17-19. Mandible rather slender, almost parallel-sided and scarcely curved, more strongly so around
the notch. Notch vestigial (never absent), formed by a dent in upper edge of mandible. Below it on the
inner side is a small thickening, forming a step in the longitudinal carina present there. Sometimes the
two structures join to form a broader step, as in the other species of the group, but smaller. Clypeal teeth
well formed and regular, the outermost pair the longest. Three centre teeth often equal, but middle one
variable. Tooth-bearing area of clypeus not very projecting, only outermost pair of teeth bent forwards,
contrasting with others. A + broad and translucent lamella connects outermost clypeal tooth with lower
orbit. Head very broad, Figs 33, 34. HW 3-5-4-0, HL 2-7-3-1, HR 1-2-1-3, EL 2-0-2-3, UIW 1-7-1:9,
MIW 1-9-2:3, LIW 1-7-1-:9, CTW 0-9-1-1, TOD 0:4-0-6, CR 1-8-2:7, MBL 2:0-2:3, MAL 0:3-0°6,
MR 3:6-5:5 (—7:3), PDL 1-9-2:3, PL 2:4—2:7, PR 1-0-1-4.
3g. BL 16-17. Clypeus with two large, sharp teeth, tooth-to-orbit margin entire (Fig. 65). Antennal
segments cylindrical, the third 0-9-1-0 long, 1-5—1-6 times as long as the first, MIW 0:15-0:20 more than
UIW. Sternite 8 with obtuse keel, tip narrowly rounded (Fig. 68). Genitalia as in Figs 93, 94. HW 3-1-3°3,
HL 2:8-2:9, HR 1-1-1-2, EL 1-9-2-0, UIW 1-6-1-7, MIW 1-8-1-9, LIW 1-4-1-5, CTW 0-30-0-32, CED
0:27-0:30, CER 1:0-1:1, TOD 0-72-0-77, CR 0:4, PDL 1-8-1-9, PL 2:6, PR 1-4. Otherwise as female.
The female is like that of mocoana, but distinguished by the mandible having a smaller notch
(sometimes vestigial) and lacking an expansion before it; and by the greater clypeal ratio. Exter-
nally, the male is extremely similar to those of cyclocephalus and moraballi, differing mainly in
its genitalia.
VARIATION. The female clypeus shows geographical variation. In the east, the teeth tend to be
sharper and the lamella less developed (Fig. 33), while in the west, the teeth tend to be more
rounded, and the lamella broader and + translucent. These are most pronounced in the Tena
specimens, in which the roundness of the teeth cannot be due to wear since two of the specimens
were reared (Fig. 34).
The propodeal dorsal rugae outside the central furrow also vary, in both sexes. They are strongly
obsolescent anteriorly in most specimens, but in those from the east they tend to be less coarse,
and moderately obsolescent also posteriorly.
DisTRIBUTION. The Amazon, from Tena to Belém; also Piaui and Tingo Maria (map, Fig. 102).
BioLoGy. Cooper found a nest at Tena, Ecuador under a leaf 1:7 m from the ground. It con-
sisted of 4 mud cells in a row, the sides in contact but without an overall covering, so that the
individual cells were plainly visible. The first cell was empty, with an emergence hole. The second
was empty but without a hole. The third contained a female wasp about to emerge. Another
150
female was later taken from cell four. This last cell (preserved in BMNH) measures about 23 mm
long x 14 mm across and has a rough surface, showing the outlines of the individual blobs of
mud composing it. Also extracted from this cell were the remains of three cockroach nymphs of
the family Blaberidae (det. J. A. Marshall). The Tena 17—19.xii female was starting to build a nest
under a leaf, about 18 ins from the ground.
MATERIAL EXAMINED. Holotype 9, EcuADpor: Napo, Tena, 9-14.xii.1971, reared (Cooper)
(BMNH).
Paratypes. ECUADOR: | 9, same data as holotype; | 9, same data except 17—19.xii.1971 (not
reared) (BMNH). Peru: 1 9, Hudnuco, Tingo Maria, 16.x.1946, 2,200 ft (Pallister) (AMNH,
New York). BRAZIL: 1 9, Para, Santarém (Wallace & Bates) (paralectotype of affinis); 1 9, Para,
Belém (both UM, Oxford); 1 9, Para, Belém, 1846 (Ghiliani) (MIZSU, Turin); 1 3, Para, Belém,
ix—x.1959 (Bianchi) (BPBM, Honolulu); 1 3, Para, Santarém (Bates) (paratype of moraballi)
(BMNH); | 9, Piaui (TM, Budapest); 1 9, Amazonas, Tefé (Bates) (ZSBS, Munich); 1 3, Ama-
zonas, Villa Nova (Bates) (det. as cyclocephalus by Richards) (BMNH); 1 9, Para, Belém,
Mocambo, 6.viii.1970 (Pimentel); 1 9, Para, Ilha das Ongas, 30.i.1977 (Overal) (both MP, Belém);
1 9, Ilha das Ongas, 22.1.1977 (Overal) (BMNH).
References
Arlé, R. 1933. Sobre a nidificagao, a biologia e os parasitos de Sceliphron (Trigonopsis) abdominalis Perty
{in both Portuguese and French]. Anais Acad. bras. Cienc. 5 : 205-212, 3 pls.
Bohart, R. M. & Menke, A. S. 1976. Sphecid wasps of the world. ix + 695 pp., frontispiece, 190 figs. Berkeley
& Los Angeles.
Cameron, P. 1888-1900. Biologia Centrali-Americana. Insects. Hymenoptera (Fossores) 2: 413 pp.,
14 pls. London.
Dalla Torre, C. G. de 1897. Catalogus Hymenoptorum hucusque descriptorum systematicus et synonymicus.
8. Fossores (Sphegidae). viiit+ 749 pp. Leipzig.
Eberhard, W. G. 1972. Altruistic behavior in a sphecid wasp: support for kin-selection theory. Science,
N.Y. 175 : 1390-1391.
—— 1974. The natural history and behavior of the wasp Trigonopsis cameronii Kohl (Sphecidae). Trans.
R. ent. Soc. Lond. 125 : 295-328, 5 figs.
Fabricius, J. C. 1804. Systema Piezatorum. 439 pp. Braunschweig.
Howes, P. G. 1969. Photographer in the rain-forests. xx+218 pp., 112 pls. Noroton, Connecticut.
Kohl, F. F. 1902. Die Hymenopterengruppen der Sphecinen. 2. Monographie der neotropischen Gattung
Podium Fabricius. Abh. zool.-bot. Ges. Wien 1: 1-101, 7 pls.
Mocsary, S. 1883. Hymenoptera nova europeae et exotica. Ertek. Term.-Tud. Kérebdl 13 (11) : 1-72.
Perty, M. in Spix, J. B. de & Martius, C. F. P. de 1833. Delectus animalium articulatorum in Brasilia
collectorum. pp. 125-224, pl. 27. Munich.
Rau, P. 1933. The jungle bees and wasps of Barro Colorado Island. 324 pp., 1 pl., 109 figs. Kirkwood.
Richards, O. W. 1937. Results of the Oxford University Expedition to British Guiana, 1929. Hymenoptera.
Sphecidae and Bembecidae. Trans. R. ent. Soc. Lond. 86 : 101-118, 1 pl.
Saussure, H. de 1867. Hymenoptera. Familien der Vespiden, Sphegiden, Pompiliden, Crabroniden und
Heterogynen. Reise der dsterreichischen Fregatte Novara um die Erde. 1857-1859. Zoologischer Teil. 2.
Hymenoptera. 156 pp., 4 pls. Vienna.
Smith, F. 1851. Descriptions of some new species of exotic Hymenoptera in the British Museum and other
collections. Ann. Mag. nat. Hist (2) 7 : 28-33.
Ee at Catalogue of hymenopterous insects in the collection of the British Museum. 497 pp., 11 pls.
ondon.
—— 1873. Descriptions of new species of fossorial Hymenoptera in the collection of the British Museum.
Ann. Mag. nat. Hist. (4) 12 : 49-59.
Vecht, J. van der 1961. Hymenoptera sphecoidea fabriciana. Zool. Verh. 48 : 1-85.
Williams, F. X. 1928. Studies in tropical wasps — their hosts and associates. Bull. Hawaiian Sug. Pirs°
Ass. Exp. Stn. 19 : 1-179, 16 figs, 33 pls.
151
abdominalis 133
affinis 141
cameronii 136
cooperi 150
cyclocephalus 144
frivaldskyi 133
grylloctonus 139
Index
Junior synonyms and other invalid names are in italics.
haemorrhoidalis 133
howesi 138
intermedia 143
menkei 133
mocoana 149
moraballi 145
neotropica 147
152
resplendens 131
richardsi 140
rufiventris 133
schunkei 142
soror 133
succinea 149
vicina 141
violascens 131
violaceus 131
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Nine genera of fungus-feeding
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J. M. Palmer & L. A. Mound
Entomology series Vol37No5 28 September 1978
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Vol 37 No 5 pp 153-215
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Nine genera of fungus-feeding Phlaeothripidae
(Thysanoptera) from the Oriental Region
J. M. Palmer & L. A. Mound
Department ent of Entomology, British Museum (Natural History), Cromwell Road, Conann
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Contents
Synopsis ; : : ‘ : : : ; ; : ; ‘ 2 133
Introduction . ‘ ’ : : : ; Fem |p)
Acknowledgements and depository abbreviations : : J , , . 154
Check-list of the species discussed in this paper . ; ; : ; , o=ri5e
Ecacanthothrips Bagnall ; : : ; : A ; : ; = ASG
Aesthesiothrips Ananthakrishnan . , : : : : : , 163
Diaphorothrips Karny . , 2 ; : ; : : . ‘ + “163
Dinothrips Bagnall : , ‘ , ‘ : ) : : } . 166
Elaphrothrips Buffa , : ; ; y ; : : ; ' pe ef
Machatothrips Bagnall . : : . ; : : ; : : . 186
Malesiathrips gen.n. . : : ' ‘ 2 . . , . 196
Mecynothrips Bagnall . : : : : : , ; ; ; . 199
Meiothrips Priesner : : : ; : ; : ‘ : . 209
References , ,- : ; ‘ : ; : : ‘ : G peeks
Index. : : : . : : : 3 ; ; A : nee Ae
Synopsis
Revisions are given of nine genera with 65 species mainly from the Oriental Region. One of these, Ecacan-
thothrips (6 species), belongs in the Phlaeothripinae; the rest are all Idolothripinae, Aesthesiothrips
(1 species), Diaphorothrips (3 species), Dinothrips (5 species), Elaphrothrips (17 Oriental species), Machato-
thrips (14 species including 3 from Africa), Mecynothrips (13 species), Meiothrips (3 species), Malesia-
thrips (3 species). A check-list is provided summarizing the nomenclatorial changes, which include one
new genus, six new species, 41 new specific synonyms and 10 new generic synonyms.
Introduction
The thrips species which feed on fungi associated with freshly dead twigs and branches are
amongst the largest and most spectacular insects in the order Thysanoptera. Many are large in
size, up to 15 mm long, and many exhibit extraordinary patterns of allometric growth and of
sexual dimorphism. However, much of the early taxonomic work on these species was carried
out before the significance of structural variation and its relationship to body size was fully
appreciated.
Recent collections by one of the authors (L. A. Mound) in Malaya and Java have facilitated
a reassessment of the species criteria which are current in such complex genera as Elaphrothrips
and Ecacanthothrips. In these two genera in particular, species are variable in structure, wide-
spread in distribution and relatively abundant. In contrast the species of Machatothrips are less
variable, more localized and relatively infrequent, and these species are peculiarly difficult to
define. It seems likely that the different patterns of speciation in these genera are related to
different patterns of activity of the individuals, but how a taxonomist should interpret such
differences in terms of names is at present very much a matter of opinion.
Bull. Br. Mus. nat. Hist. (Ent.) 37 (5): 153-215 Issued 28 September 1978
153
The revisions given here concern one genus of Phlaeothripinae and eight genera of Idolothri-
pinae; these include all the common species of large Thysanoptera which are found in this habitat
in the Oriental region. It is intended to re-examine the generic classification of the Idolothripinae
elsewhere, but such a study must be built on a sound knowledge of the variation within and
between species. For example, Mecynothrips is used here to include a group of 12 species which
have been divided previously between five genera and two subgenera. A study of the variation
within and between these species has indicated that large and small males of certain species would
be placed in separate genera using traditional criteria, but that it is almost impossible to distinguish
between the females at species level. This process of reassessment of taxonomic criteria must
needs involve not only recently collected specimens but also the type-material of earlier described
species. In this paper all the material which has been studied is detailed, and in the few instances
where type-specimens were not available the species concerned are not dealt with in the keys
but are merely listed in the text. This approach probably explains the difference in interpretation
of certain names, particularly in Elaphrothrips, between the present work and that of other recent
authors.
Acknowledgements and depository abbreviations
This revisionary study has been carried out with the cooperation of numerous colleagues who are
responsible for collections in other institutes and museums. The authors are very grateful to the
following for loans of material; depository abbreviations used in the text are given in parentheses.
Professor T. N. Ananthakrishnan, Director, Zoological Survey of India, Calcutta (TNA); Paul
H. Arnaud, California Academy of Sciences, San Francisco (CAS); E. Dahms, Queensland
Museum, Brisbane (QM); C. F. Jacot-Guillarmod, Albany Museum, Grahamstown (JG); A.
Kaltenbach, Naturhistorisches Museum, Wien (NM); I. Kudo, Shizuoka Seikogakuin, Oshika,
Japan; Madame D. Mathilde-Ferrero, Muséum National d’Histoire Naturelle, Paris (MNHN);
Shuji Okajima, Tokyo University of Agriculture (SO, Tokyo); Kellie O’Neill, U.S.D.A., Belts-
ville, Maryland (USNM); M. Suwa, Faculty of Agriculture, Hokkaido University, Sapporo
(HUS); R. zur Strassen, Senckenberg Museum, Frankfurt (SMF).
The Stereoscan scanning electron micrographs (Figs 22, 23) were provided by our colleague
Brian R. Pitkin. The photomicrographs (Figs 1-4) were provided by the Photographic Unit of
the British Museum (Natural History) (later referred to as BMNH in the text).
Check-list of the species discussed in this paper
Subfamily PHLAEOTHRIPINAE Uzel erythrinus Ananthakrishnan, 1956
ECACANTHOTHRIPS Bagnall, 1909 ch gg
etcheri Ramakrishna, 1
Ormothrips Buffa, 1909 as Girault, sb ae ae
matsumurai Ishida, 1934 syn. n.
priesneri Hood, 1935
ramakrishnai Ananthakrishnan, 1952
andrei sp. n.
coniger Priesner, 1930
inarmatus Kurosawa, 1932
piceae Ishida, 1936 syn. n.
leai Moulton, 1947
spinipes (Bagnall, 1908)
bagnalli Priesner, 1930
guineaensis Moulton, 1947
inermis (Buffa, 1909)
tibialis (Ashmead, 1905) comb. n.
bryanti Bagnall, 1915
coxalis Bagnall, 1915
coxalis consanguineus Priesner, 1930
coxalis formosensis Takahashi, 1936 syn. n.
coxalis philippinensis Priesner, 1930 syn. n.
crassiceps Karny, 1913 syn. n.
sanguineus (Bagnall, 1908) syn. n.
steinskyi (Schmutz, 1913)
Subfamily IDOLOTHRIPINAE Bagnall
AESTHESIOTHRIPS Ananthakrishnan, 1961
jatrophae Ananthakrishnan, 1961
DIAPHOROTHRIPS Karny, 1920
D. (Cnemidothrips) Priesner, 1940
clavipes Priesner, 1940
hamipes Karny, 1923
unguipes Karny, 1920
spinosus Ananthakrishnan, 1959
thevetii Ananthakrishnan, 1957 syn. n.
154
DINOTHRIPS Bagnall, 1908
Paxillothrips Ananthakrishnan, 1961 syn. n.
juglandis Moulton, 1933
longicauda (Ananthakrishnan, 1961) comb. n.
monodon Karny, 1920
spinosus (Schmutz, 1913) comb. n.
affinis Bagnall, 1915 syn. n.
anodon Karny, 1923 syn. n.
celebensis Bagnall, 1934 syn. n.
crassiceps (Bagnall, 1921) syn. n.
gardneri Moulton, 1928 syn. n.
jacobsoni Karny, 1921 syn. n.
kemneri Karny, 1923 syn. n.
malloti Moulton, 1933 syn. n.
sumatrensis Bagnall, 1908
fulmeki Priesner, 1959 syn. n.
furcifer Schmutz, 1913
ELAPHROTHRIPS Buffa, 1909
Dicaiothrips Buffa, 1909
E. (Elaphoxothrips) Bagnall, 1932
E. (Cradothrips) Ananthakrishnan, 1973
amoenus Priesner, 1935
athletes (Karny, 1923)
bakeri (Karny, 1920)
bakeri depokensis Priesner, 1935 syn. n.
imitator Priesner, 1935 syn. n.
mentaweiensis Priesner, 1929 syn. n.
coreanus Woo, 1974
curvipes Priesner, 1929
karnyi Priesner, 1935 syn. n.
secus Ananthakrishnan, 1973 syn. n.
denticollis (Bagnall, 1909)
beesoni Ramakrishna Ayyar, 1934 syn. n.
mucronatus Priesner, 1935
productus Priesner, 1935 syn. n.
productus obscuricornis Priesner, 1935 syn. n.
sumbanus Priesner, 1935 syn. n.
fulmeki Priesner, 1935
greeni (Bagnall, 1914)
bouvieri (Vuillet, 1914)
micidus Ananthakrishnan, 1973 syn. n.
insignis Ananthakrishnan, 1973
insularis Priesner, 1928
jacobsoni Priesner, 1935
mahensis (Bagnall, 1921)
hystrix (Bagnall, 1921)
rex (Bagnall, 1921)
malayensis (Bagnall, 1909)
bruneitarsis (Schmutz, 1913) syn. n.
bruneitarsis levis (Schmutz, 1913) syn. n.
coronatus Bagnall, 1934 syn. n.
notabilis Ananthakrishnan, 1973
procer (Schmutz, 1913)
achaetus Bagnall, 1934
approximatus Bagnall, 1934
chandana Ramakrishna Ayyar, 1934
dallatorensis (Schmutz, 1913)
eranthemi Seshadri & Ananthakrishnan, 1954
novus (Schmutz, 1913)
proximus (Bagnall, 1914)
sensitivus Priesner, 1929
seychellensis (Bagnall, 1921)
spiniceps Bagnall, 1932
clarispinis Priesner, 1935 syn. n.
graveleyi Bagnall, 1934 syn. n.
MACHATOTHRIPS Bagnall, 1908
Adiaphorothrips Bagnall, 1909
Cnestrothrips Priesner, 1939 syn. n.
antennatus (Bagnall, 1915)
dammermani (Priesner, 1939) syn. n.
artocarpi Moulton, 1928
biuncinatus Bagnall, 1908
montanus Priesner, 1932 syn. n.
simplex (Bagnall, 1909)
braueri Karny, 1912
multidens Bagnall, 1934
paucidens Bagnall, 1934
paucidens bicolorisetosus Bagnall, 1934
braueri karnyi Priesner, 1932
celosia Moulton, 1928
corticosus Ananthakrishnan, 1972
decorus sp. n.
diabolus Priesner, 1928
haplodon Karny, 1925
braueri buffai Karny, 1925
braueri spatiata Priesner, 1932 syn. n.
simplicidens Bagnall, 1934
heveae Karny, 1921
indicus Ananthakrishnan & Jagadish, 1970
lentus sp. n.
quadrudentatus Moulton, 1947
silvaticus Ananthakrishnan, 1972
MALESITATHRIPS gen. n.
guamensis sp. n.
malayensis sp. n.
solomoni (Mound, 1970) comb. n.
MECYNOTHRIPS Bagnall, 1908
Acrothrips Karny, 1921 syn. n.
Dracothrips Bagnall, 1914 syn. n.
Kleothrips Schmutz, 1913 syn. n.
K. (Akleothrips) Priesner, 1935 syn. n.
K. (Synkleothrips) Priesner, 1935 syn. n.
Phoxothrips Karny, 1913 syn. n.
acanthus (Hood, 1918) comb. n.
gargantua (Girault, 1926)
giganteus (Girault, 1926)
sorex (Karny, 1921)
atratus (Hood, 1919) comb. n.
zuluensis (Jacot-Guillarmod, 1939) syn. n.
goliath (Priesner, 1935) comb. n.
hardyi (Priesner, 1928) comb. n.
kanoi (Takahashi, 1937) comb. n.
karimonensis (Priesner, 1935) comb. n.
karimonensis parvidens (Priesner, 1935)
syn. n.
Kraussi sp. n.
lacerta (Priesner, 1935) comb. n.
innocens (Priesner, 1935) syn. n.
priesneri Mound, 1971 bagnalli imbecilla Priesner, 1935 syn. n.
minor Mound, 1971 syn. n. magnus Girault, 1929
pugilator (Karny, 1913) comb. n. MEIOTHRIPS Priesner, 1929
takahashi (Priesner, 1935) M. (Telothrips) Kudo & Ananthakrishnan,
simplex Bagnall, 1912 comb. rev. 1974 (preocc.) syn. n.
agama (Priesner, 1935) M. (Aculeathrips) Kudo, 1975 syn. n.
ceylonicus (Bagnall, 1914) annulipes (Bagnall, 1914)
gigans (Schmutz, 1913) annulatus Priesner, 1929 syn. n.
snodgrassi Hood, 1952 menoni Ananthakrishnan, 1964
wallacei Bagnall, 1908 nepalensis Kudo & Ananthakrishnan, 1974
bagnalli Priesner, 1935
ECACANTHOTHRIPS Bagnall
Ecacanthothrips Bagnall, 1909a: 348. Type-species: Acanthothrips sanguineus Bagnall, by monotypy
(here regarded as a synonym of Jdolothrips tibialis Ashmead).
Ormothrips Buffa, 1909 : 166. Type-species: Ormothrips inermis Buffa, by monotypy (here regarded as a
synonym of Phloeothrips spinipes Bagnall). [synonymized by Bagnall, 1911 : 464.]
Macropterous, sexually dimorphic Phlaeothripinae. Antennae 8-segmented; segment IV with 4 sense
cones, III with enlarged and usually supernumerary sense cones. Head with large eyes, cheeks with at
least one pair of setae on small tubercles; stylets retracted almost to eyes, close together in middle of head;
mouth cone pointed. Pronotum transverse, epimeral sutures complete; all major setae developed, antero-
angulars frequently elongate in 3; praepectus absent. Fore tarsus usually with a tooth; fore femur of 3
with a pair of apical tubercles; both sexes sometimes with a median fore femoral tubercle. Mesonotal
lateral setae elongate; metanotum reticulate. Fore wing constricted medially, duplicated cilia present.
Pelta bell-shaped, rarely almost triangular; tergites II-VII with supernumerary wing retaining setae
laterally in addition to the two main pairs; sternite VIII of 3 without a glandular area.
Although 23 species-group names have been placed in this genus only six species are here accepted
as valid. The concept of the genus is usually based on the polymorphic species tibialis Ashmead
(=sanguineus Bagnall), and involves two obvious characters — the large number of enlarged
sense cones on the third antennal segment, and the well-developed tubercle on the inner margin
of the fore femur. However, only five of the six species here recognized in the genus have the
first of these two characters, and only two of the species have the second character. One species,
described below as andrei, has a well-developed fore femoral tubercle but has only four enlarged
sense cones on the third antennal segment (Figs 5-8). This species is thus intermediate between
the genera Ecacanthothrips and Hoplandrothrips. The alternative combination of characters
involving four enlarged sense cones and unarmed fore femora is found in Hoplandrothrips
flavipes Bagnall, and this species could well be treated in Ecacanthothrips as a close relative of
andrei. However, the characteristic of enlarged sense cones on the third antennal segment is not
in itself clear-cut in other undescribed species from Malaya and Japan. As a result, Ecacantho-
thrips could well be regarded as an Oriental species-group derived from the world-wide genus
Hoplandrothrips.
Key to species
1 Fore femur of 2 and ¢ with a stout median tubercle on inner margin, in very small 9 this
tubercle is scarcely visible; ¢ with a pair of tubercles at apex of fore femur (Figs 6-8). : P.
— Fore femur of 2? and ¢ with inner margin smooth or bearing a series of small tuberculate setae;
3 with a pair of tubercles at apex of fore femur . 3
2 Antennal segment III with at least 10 sense cones; pelta triangular; mid and hind femur with
3 stout capitate setae on anterior margin [Widespread] . . tibialis (p. 161)
— Antennal segment III with 4 sense cones (rarely 5, 3 or 2) (Fig. 5); pelta broadly bell-shaped ;
anterior margin of mid and hind femur with one stout capitate seta [Malaya] . andrei (p. 158)
3 Anterior margin of mid and hind femur with a row of 5 or more stout setae with blunt or
capitate apices; pelta broadly triangular; inner margin of fore femur with a row of short
stout setae on small tubercles [New Guinea, Solomon Is] ; y p . Spinipes (p. 161)
156
Figs 1-4 Ecacanthothrips tibialis, (1, 2) 3; (3, 4) 2.
157
— Anterior margin of mid and hind femur with a row of 10 or more fine setae with acute apices;
pelta bell-shaped; inner margin of fore femur smooth . 4
4 Setae B, on tergite IX 1-1 times as long as tube; pronotum with well- developed closely spaced
transverse lines of sculpture; sculptured reticles of metanotum and pelta with internal linear
markings; posterolateral areas of tergites II-VII with numerous very stout dentate micro-
trichia; antennal segment III with more than 20 black sense cones which occupy about half
of the ventral surface of the segment [Borneo] : . coniger (p. 160)
— Setae B, on tergite IX less than 0-9 times as long as tube; pronotal sculpture weak, without
transverse lines; sculptured reticles without internal markings; tergal microtrichia weak,
ciliate or absent: antennal segment III with 15 sense cones or less . ‘ 2.
5 Fore wing pale or weakly shaded, but dark brown around sub-basal setae [Japan] inarmatus (p. 160)
Fore wing pale or shaded, but not dark at base except on wing scale [Japan to Solomon Is]
leai (p. 160)
Ecacanthothrips andrei sp. n.
(Figs 5-8)
2 macroptera. Colour brown with some red hypodermal pigment; mid and hind tibiae and apices of femora
clear yellow, fore tibiae yellow with faint brown markings; antennal segment III brownish yellow,
succeeding segments increasingly dark, IV—VI light brown with yellowish pedicels; major setae of head
and thorax dark, of wings and abdomen pale; wings light brown, paler at base, at median constriction
and a little paler at apex.
Head reticulate dorsally but reticulation weak medially ; cheeks slightly swollen, with one pair of major
setae in posterior third. Antennal segment III with 4 sense cones ventrally, less commonly with 5, or 3,
or only 2, segment IV with 4 sense cones (Fig. 5). Pronotum sculptured medially; fore femur with hooked
median tubercle (Fig. 8); fore tibia with 4 to 5 small tubercles on inner margin; fore tarsal tooth variable
in size. Mid and hind femora with 1 stout seta on anterior margin. Pelta broadly bell-shaped; tergites
IIJ-VII with a transverse row of 4 setae lateral to wing retaining setae; lateral areas with microtrichia on
the lines of sculpture.
Measurements (holotype 2 in um). Body, length 3100. Head, length 330; maximum width 230; post-
ocular setae 120; basal cheek setae 26. Pronotum, length 195; median width 370; major setae, antero-
marginal 75, anteroangular 90, mid-lateral 80, epimera! 90, posteroangular 130. Width of prothorax
across coxae 550. Mesonotal lateral setae 50. Fore wing, length 1150; distal width 80; sub-basal setae 70,
80, 80; number of duplicated cilia 15. Tergite IX setae B, 130. Tube, length 175. Antennal segments
III-VIII length, 90, 115, 100, 68, 55, 40.
3 macroptera. Colour similar to 2. Body size variable, large 3 with elongate head bearing 3 pairs of
stout cheek setae, elongate femoral tubercle (Fig. 7), long pointed coxae and large fore tarsal tooth;
small 3 with short head bearing weak cheek setae, very small fore femoral tubercle and fore tarsal tooth
(Fig. 6), coxae normal and rounded.
Measurements (largest and smallest ¢ paratypes). Body, length 2800 (1650). Head, length 350 (225);
maximum width 190 (165); postocular setae 165 (70); basal cheek setae 35 (16). Pronotum, length 200
(97); median width 320 (210); major setae, anteromarginal 210 (42), anteroangular 115 (50), mid-lateral
115 (38), epimeral 100 (50), posteroangular 130 (65). Width of prothorax across coxae 550 (275). Fore
wing, length 1000 (700); number of duplicated cilia 14 (7). Tube, length 160 (115). Antennal segments,
III 90 (75); IV 100 (70).
SPECIMENS STUDIED.
Holotype 2, Malaya: Gombak Field Station, 30 km east of Kuala Lumpur, on dead branches, 5.x.1973
(L. A. Mound) (BMNH).
Paratypes. Malaya: 7 9, 7 ¢ with same locality and collector as holotype, 29.ix.1973—7.x.1973; Kuala
Lumpur, 3 2, 6 ¢ on partly dead branch, 27.xii.1969 (R. & F. Andre) (BMNH). Singapore: Bukit Timah
Reserve, 1 2 on dead branch, 3.xi.1973 (L. A. Mound) (BMNH).
The head, pronotum and fore legs of the male of andrei are very similar to those of tibialis,
moreover they show the same range of structural variation (Figs 6-8). However, large males have
the pronotal anteromarginal setae much longer than the anteroangular setae instead of reduced
as in tibialis, and the second pair of postocular setae are short and pointed, not elongate and
expanded. The females are much less variable than the males. Most of the specimens studied have
158
<
Figs 5-15 5-8, Ecacanthothrips andrei, (5) 2 right antenna; (6, 7) 3 fore leg; (8) ? fore leg. 9-11,
Aesthesiothrips jatrophae, (9) head and pronotum; (10) tergite III; (11) 2 fore tibia and tarsus.
12 Diaphorothrips clavipes °, fore tibia and tarsus. 13, 14, peltas, (13) A. jatrophae; (14) D. clavipes.
15, A. jatrophae, right antenna.
159
4 sense cones on the third antennal segment, but several individuals have one or both antennae
with only 3 or 2 sense cones on this segment, and one specimen has 5 sense cones on this segment
on one of its antennae. This variation is not related to body size, but when the number of sense
cones on a segment is reduced then their own individual size is increased. As discussed in the
introduction, andrei is intermediate in structure between Ecacanthothrips and Hoplandrothrips.
Ecacanthothrips coniger Priesner
Ecacanthothrips coniger Priesner, 1930 : 363-364. Syntypes 2 9, BoRNEO (SMF) [1 2 examined].
The only specimen of this species which has been studied is a large female. This specimen is easily
recognized from the characters given in the key, but at least some.of these characters are probably
related to the large body size. If a small female of coniger is ever collected it will probably be
rather similar to the other members of the inarmatus-group.
SPECIMEN STUDIED.
Borneo: Midden, | 2 ‘paratype’, 16.viii.1925 (H. C. Siebers 33) (SMF).
Ecacanthothrips inarmatus Kurosawa
Ecacanthothrips inarmatus Kurosawa, 1932 : 238-242. Holotype 2, JAPAN (Nat. Inst. Agric. Sci. Tokyo)
[examined].
Ecacanthothrips piceae Ishida, 1936: 154-156. Holotype 3, JAPAN (? lost) [not examined]. Syn. n.
This species appears to be confined to the Japanese mainland. It is similar to, but considerably
larger than, /eai from Malaya and Java. The pattern of intraspecific variation in the inarmatus
species-group requires further study based on new collections. Haga (1972) states that piceae
may be the male form of inarmatus. The unique holotype of piceae is not in the Ishida collection
at Sapporo and there is nothing in the description to distinguish it from inarmatus.
The smallest specimen listed below has a head rather similar to /eai, but the other specimens
are larger with a relatively long head. The pronotal anteromarginal setae of the males are elongate,
about two-thirds as long as the anteroangulars, and the males have 2 pairs of long stout setae
laterally on the posterior half of the head. The colour of the legs and antennal segments appears
to be variable from yellow to brown, but even in the palest specimen the tibiae are dark.
SPECIMENS STUDIED.
Japan: Sizuoka, lida-mura, holotype 2 of inarmatus in hole of citrus tree, 24.xi.1930 (7. Noguti) (SO,
Tokyo); Kanagawa Prefecture, Atsugi, 1 3, 1 2 on dead leaves, 18.iii.1975; Kawada, Mikura-jima L.,
2 2 (2 BMNH) on dead leaves, 9.vi.1973 (SO, Tokyo), 1 9, 11.vi.1973 (BMNH); Ohdaru spa, 1 ¢ on
dead leaves, 27.vi.1972 (SO, Tokyo).
Ecacanthothrips leai Moulton
Ecacanthothrips leai Moulton, 1947 : 176. Holotype 2, MALAYA (CAS) [examined].
This species was described from a single damaged female, but further specimens of both sexes
have now been collected in both Malaya and Java. Moreover two females from New Guinea,
referred to as spinipes by Mound (1970), are here regarded as /eai as well as two specimens from
the Ryuku Islands and four specimens from the Solomon Is. In Malaya /eai was collected with
both tibialis and andrei, and tibialis is also known from the Ryuku Is.
The differences between /eai and spinipes are referred to under the latter species. Unfortunately
all of the differences could be a function of size, that is /eai may represent a small form of spinipes.
Similarly the differences between /eai and inarmatus are of the same order as the geographical
variation in colour noted in tibialis. The male and female listed below from Amami-ohshima
(Ryuku Is) are larger than the /eai specimens from Malaya and Java with the head relatively
long and slender and the pronotal anteromarginal setae of the male as long as the anteroangular
160
setae. Major males of /eai from Malaya have the anteromarginal setae reduced. One of the females
listed below from New Guinea has brown hind tibiae, whereas all the other specimens of /eai
have yellow tibiae. Delimitation of species in the inarmatus-group is still far from clear, and the
description of further species needs to be avoided until more material is collected.
SPECIMENS STUDIED.
Malaya: Kuala Lumpur, 2 holotype (A. M. Lea) (CAS); Kuala Lumpur, 11 2, 2 ¢ on dead branches
xii.1969 (R. & F. Andre) (BMNH); Gombak, 30 km east of Kuala Lumpur, 5 2, 2 3 on dead wood,
ix—x.1973 (L. A. Mound) (BMNH). Java: Tjibodas Botanic Garden, 5 2, 2 d on dead leafy twigs, x.1973
(L. A. Mound) (BMNH). New Guinea: Wau, Bulolo Gorge, 900 m, 1 2 from bark of fallen log, 4.vi.1968
(R. Rice); Aiyura, 5400 ft [1800 m], 1 2 on grasses, 22.ii.1962 (J. H. Barrett) (BMNH). Solomon Is:
Rendova, 2 3, 2 2 on dead leaves, xi.1972 (Krauss) (BMNH). Japan: Amami-ohshima Is, Nishinakama,
1 2, 1 3, 30.vi.1972 (M. Kuboki) (SO, Tokyo).
Ecacanthothrips spinipes (Bagnall)
Phloeothrips spinipes Bagnall, 19085 : 195-196. Holotype 9, NEw GuINEA (BMNH) [examined].
Ecacanthothrips spinipes (Bagnall) Mound, 1968 : 90.
Ormothrips inermis Buffa, 1909: 168. Holotype ?, NEw GuINEA (? lost) [not examined]. [Synonymized
by Mound, 1970: 96.]
Ecacanthothrips bagnalli Priesner, 1930: 364-365. Holotype 3, Kr1 Is (SMF) [examined]. [Synonymized
by Mound, 1970: 96.]
Ecacanthothrips guineaensis Moulton, 1947: 176-177. Holotype 2, New GuineA (CAS) [examined].
[Synonymized by Mound, 1970: 96.]
Mound (1970) listed material of this species from the Solomon Islands and New Guinea and
discussed variation in the shape of the pelta which appeared to be dependent on body size.
However, the two small females from New Guinea (1 9 Aiyura, 1 2 Wau) which have a bell-
shaped pelta also lack capitate setae on the mid and hind femora. These specimens are here
regarded as /eai despite the fact that one of them was collected with 11 9, 1 3 of spinipes. Unfor-
tunately all of the known spinipes specimens are considerably larger than any of the known
leai specimens. Therefore it is impossible to be certain that the obvious differences in head shape,
pelta and femoral setae are not a function of size rather than real species differences. In contrast
to /eai the inner margin of the fore femur bears a series of small tuberculate setae, the mid and
hind femora bear a row of stout capitate setae, the head is relatively short and narrowed to the
base with three or more pairs of stout tuberculate setae in the female and four or more pairs of
long stout setae in the male. The colour of the tibiae and third antennal segment appears to be
variable. No further material has been studied apart from that listed by Mound (1970: 97)
from New Guinea and the Solomon Is.
Ecacanthothrips tibialis (Ashmead) comb. n.
(Figs 1-4)
Idolothrips tibialis Ashmead, 1905 : 20. Holotype 2, PHILIPPINES (USNM) [examined].
Acanthothrips sanguineus Bagnall, 1908a : 362. Holotype 9, NEw GUINEA (BMNH) [examined]. Syn. n.
Ecacanthothrips sanguineus (Bagnall) Bagnall, 1909a : 348.
Ormothrips sanguineus (Bagnall) Buffa, 1909 : 166.
Ecacanthothrips crassiceps Karny, 1913 : 134-136. Holotype ? 29, NEw Guinea (? lost) [not examined].
Syn. n.
Ormothrips Steinskyi Schmutz, 1913 : 1028-1030. Syntypes 3, 2, Ski LANKA (Ceylon) (NM) [2 9 examined].
[Synonymized with sanguineus by Ananthakrishnan, 19615 : 275.]
Ecacanthothrips bryanti Bagnall, 1915a : 320-321. Lectotype $, SARAWAK (BMNH) [examined]. [Synony-
mized with sanguineus by Ananthakrishnan, 19615: 275.]
Ecacanthothrips coxalis Bagnall, 19155 : 597. Lectotype 3, SARAWAK (BMNH) [examined]. [Synonymized
with sanguineus by Ananthakrishnan, 19615: 275.]
Ecacanthothrips flavipes Bagnall, 19155: 597. Lectotype ¢, SARAWAK (BMNH) [examined]. [Synony-
mized with sanguineus by Ananthakrishnan, 1961 : 275.]
161
Ecacanthothrips coxalis var. consanguineus Priesner, 1930 : 367-368. Syntypes 3, 9, MALAYA & BORNEO
(SMF & BMNH) [examined]. [Synonymized with sanguineus by Mound, 1968 : 89.]
Ecacanthothrips coxalis var. philippinensis Priesner, 1930: 368. Syntypes ?, PHILIPPINES (? SMF) [not
examined]. Syn. n.
Ecacanthothrips io Girault, 1930: 1. Holotype 3, AUSTRALIA (QM) [examined]. Syn. n.
Ecacanthothrips matsumurai Ishida, 1934: 149-151. Holotype 3, TAIWAN (? lost) [not examined]. Syn. n.
Ecacanthothrips fletcheri Ramakrishna, 1934: 495-496. Holotype 9, INDIA (? TNA) [not examined].
[Synonymized with sanguineus by Ananthakrishnan, 1961b: 275.]
Ecacanthothrips priesneri Hood, 1935: 196-199. Holotype 2, TANZANIA (USNM) [examined]. [Synony-
mized with sanguineus by Ananthakrishnan, 19615: 275.]
Ecacanthothrips coxalis var. formosensis Takahashi, 1936 : 454-456. Holotype 3, TAIWAN (? Dept. Agric.
Res. Taiwan) [not examined]. Syn. n.
Ecacanthothrips ramakrishnai Ananthakrishnan, 1952 : 38-41. Holotype 3, INDIA (TNA) [not examined].
[Synonymized with sanguineus by Ananthakrishnan, 19616: 275.]
Ecacanthothrips erythrinus Ananthakrishnan, 1956: 134-136. Holotype 3, INDIA (TNA) [not examined].
[Synonymized with sanguineus by Ananthakrishnan, 19615: 275.]
Ashmead described tibialis from a single specimen which at that time was mounted dry on a
card point. This female was subsequently placed in balsam on a slide and is a typical member of
the species usually known as sanguineus. Karny described crassiceps from a single specimen from
New Guinea. This specimen is not available but the description defines satisfactorily an extreme
gynaecoid male of tibialis, particularly when it is remembered that both Karny and Priesner
measured ‘head length’ from the anterior margin of the eyes instead of the extreme anterior
margin of the head. The specimen described as io by Girault is a typical male of tibialis. More-
over, although type-specimens have not been studied there is nothing in the descriptions of
Philippinensis, matsumurai nor formosensis to distinguish these from the species tibialis which is
known to be common in both the Philippines and also Formosa.
This species is exceptionally variable. Structural variation associated with allometric growth
is found throughout the geographical range of the species and has been discussed and illustrated
by Hood (1935) and Ananthakrishnan (1961). Variation in colour of the tibiae and also the
median antennal segments does not seem to be associated with body size, but there is a tendency
for specimens from the western end of the range to have darker appendages than specimens
from the eastern end. Thus the specimens from the Philippines and many of those from Borneo
have the pedicels of antennal segments V and VI yellow, and the hind tibiae vary from yellow
washed with brown medially to dark brown with paler ends. Most of the specimens studied from
Malaya, Sri Lanka, India, Mauritius and Tanzania have the antennal pedicels brown and the
hind tibiae frequently black. However, the specimens listed below from Japan and Taiwan are
also dark. These colour variations, together with the structural variation associated with allo-
metric growth patterns (Figs 1-4), account for the large number of names by which this species
has been known.
SPECIMENS STUDIED.
Japan: Okinawa-jima Is, 3 g, 1 2 (SO, Tokyo; 1 2 BMNH); Amami-ohshima Is, 1 9, vii.1972 (SO,
Tokyo; 1 § BMNH). Taiwan: Nansanchi, 2 9, 2 3, vi.1972 (SO, Tokyo); Heito, 1 2 from bark of Arto-
carpus, 25.i1i.1933 (M. Ishida 1685) (HUS) [det. sanguineus by Haga, 1972, and labelled ‘M. Ishida’s E.
matsumurai’]. China: Chengtu, | 9, viii.1930 (BMNH). Vietnam: Tonkin, 4 2, 3 ¢ (BMNH). Philippines:
Manilla; holotype 2 of tibialis (Robt Brown) (USNM); 17 9, 8 3, vii.1931 (BMNH). Borneo: Midden,
1 3, 1 2 syntypes of consanguineus, viii.1925; Sarawak, Mt Matang, 3 lectotypes of bryanti, coxalis and
flavipes, xii.1913-ii.1914; Hii Siew, 3 2, 2 ¢ on Cacao (BMNH). New Guinea: Dorey, holotype 9 of
sanguineus; Wau, 2 2, 2 3, vi.1968 (BMNH). Australia: Queensland, holotype ¢ of io (QM). Indonesia:
Isle of Nias, .1 2, 7 3; Java, Bogor, 31 2, 21 3 10 instar II, x.1973 (Mound) (BMNH). Singapore: Bukit
Timor, 7 9, 12 3, xi.1973-i.1974 (BMNH). Malaya: near Kuala Lumpur, 48 9, 41 3, x.1973; Ringlet,
Cameron Highlands, 4 2, 5 3, x.1973 (Mound); Trengannu, 9 9, 2 3, iii.1974 (BMNH). India: Madras,
4 2, 5 3 (Ananthakrishnan) (BMNH). Sri Lanka: Peradeniya, 7 9, 6 3 (Green & Rutherford) (BMNH);
2 2 syntypes of Steinskyi on bark, 24-25.v.1902 (Uze/ 193, 194) (NM). Mauritius: 1 9, v.1970; 1 3, xii.1971;
1 3, vi.1971 (BMNH). Rodrigues: 1 ¢ (BMNH). Tanzania: Amani, holotype 2 of priesneri, i.1928 (USNM).
162
AESTHESIOTHRIPS Ananthakrishnan
Aesthesiothrips Ananthakrishnan, 1961a: 253. Type-species: Aesthesiothrips jatrophae Ananthakrishnan,
by monotypy.
Large, dark, macropterous species of Cryptothripini. Head elongate, eyes relatively small but slightly
prolonged ventrally; stylets retracted to eyes, close together in middle of head; postocellar setae reaching
to apex of antennal segment I; mouth cone broadly rounded. Antennae 8-segmented, III about two-
thirds as long as IV; sense cones on III and IV longer than their segment, 2 on III, 4 on IV. Pronotum
transverse, epimeral sutures complete, setae in normal position; posteroangular setae scarcely two-thirds
as long as epimerals; praepectus present. Both sexes with stout fore tarsal tooth, apex of fore tibia
flattened and bearing a small tubercle. Metanotum weakly reticulate, median setae less than 50 um long.
Fore wing relatively slender, sub-basal setae not elongate. Pelta irregularly triangular (Fig. 13); tergites
II-VII with 1 pair of wing-retaining setae, and laterally with a transverse row of 2 to 5 short setae; tergite
IX setae almost as long as tube; 2 with tube slightly longer than head, margins weakly convex; 3 with tube
shorter than head.
This genus was erected for a single species described from two damaged females. The original
description referred to the elongate sense cones on the third and fourth antennal segments, but
did not mention the remarkable maxillary stylets and long postocellar setae (Fig. 9). The form of
the head is similar to Phaulothrips Hood from Australia (Mound, 1974), but the structure of the
antennae and sense cones is quite different (Fig. 15). Celidothrips Priesner is also related but has
long postocular setae and short sense cones, and one ommatidium is isolated behind each eye on
the cheeks.
Aesthesiothrips jatrophae Ananthakrishnan
(Figs 9-11, 13, 15)
Aesthesiothrips jatrophae Ananthakrishnan, 1961a : 253-254. Holotype 9, INDIA (TNA) [examined].
This species was based on two damaged females collected in northern India. The specimens listed
below from Malaya cannot be distinguished at present from jatrophae, although the holotype has
fewer setae laterally on each tergite. The males are very similar to the females although a little
smaller with a shorter tube. However, the length of the tube is apparently subject to allometric
growth in this species. The short third antennal segment is mainly yellow in contrast to the dark
brown of the rest of the body.
SPECIMENS STUDIED.
India: Dehra Dun, Nakronda, holotype 2 on Jatropha curcus fruits, 23.iii.1934 (Bahadur) (TNA). Malaya:
Kuala Lumpur, 13 9, 4 ¢ on dead branch, 26.xii.1969 (R. G. & F. Andre) (BMNH).
DIAPHOROTHRIPS Karny
Diaphorothrips Karny, 1920a : 186. Type-species: Diaphorothrips unguipes Karny, by monotypy.
Diaphorothrips (Cnemidothrips) Priesner, 1940: 403. Type-species: Diaphorothrips hamipes Karny, by
original designation.
Medium sized, dark brown members of Cryptothripini. Head rectangular, longer than broad; cheeks
with at least 8 pairs of setae; 1 pair of ocellar setae frequently elongate; maxillary stylets retracted almost
to postocular setae, not close together in midline; mouth cone appears pointed. Antennae 8-segmented,
VIII lanceolate in shape; 2 sense cones on III, 5 on IV. Pronotum with anterior margin thickened, at
least in large specimens, anteromarginal setae reduced; both sexes with fore tarsal tooth well developed
and fore tibia with subapical projection (Fig. 12); praepectus well developed, mesopraesternum boat-
shaped. Fore wings broad with numerous duplicated cilia. Pelta hat-shaped with lateral margins curving
forwards away from tergite II (Fig. 14). Tergites with 1 pair of wing-retaining setae; setae on IX shorter
than tube; tube heavy, margins slightly convex.
The three species now recognized in Diaphorothrips are unguipes from Sri Lanka and southern
India, hamipes from Java, Sumatra and Malaya, and clavipes from both Singapore and the adjoin-
ing Riau Is. However, none of these species appears to occur in large numbers, each having been
163
collected infrequently from dead twigs or under bark. The genus is a typical member of the
Cryptothripini and is intermediate in structure between certain species of Scotothrips or Dichaeto-
thrips and the Oriental genus Uredothrips. The subgenus Cnemidothrips does not seem to be a
useful category in view of the many similarities between the three species indicated below.
Key to species
1 Antennal segment III yellow to yellowish brown in basal half, relatively short and conical, less
than 2:2 times as long as maximum width; relatively small aie with tube 1-2 times as long
as head [Malaya to Sumatra] . . hamipes (p. 164)
— Antennal segment III dark brown with extreme base pale, more than 2: s times as long as wide;
relatively large species with tube 1-4 times as long as head : 2
2 Postocellar setae more than 3-0 times as long as anteocellar setae; cheeks with 10-12 pairs of
relatively weak setae, longest cheek seta less than 26 um; fore wing clear or weakly shaded
in distal half [India, Sri Lanka] . ‘ . . unguipes (p. 164)
— Postocellar setae less than 0-8 times as long as anteocellar setae; cheeks with 8 pairs of stout dark
setae, longest cheek seta more than 30 um; fore wing deeply shaded in distal half [Riau Is,
Singapore] : ; ; : 4 . , ; : : : . Clavipes (p. 164)
Diaphorothrips clavipes Priesner
(Figs 12-14)
Diaphorothrips (Cnemidothrips) clavipes Priesner, 1940 : 403-405. Holotype 2, R1Au Is (SMF) [examined].
The cheek setae of this species are larger and darker than in the other two species. The antennae
are similar to unguipes but the ocellar setae are similar to hamipes.
SPECIMENS STUDIED.
Riau (Riouw) Is: Doerian, 2 holotype, xi.1923 (Dammerman) (SMF). Singapore: 3 2 on dead twigs,
4.xi.1973 (L. A. Mound) (BMNH).
Diaphorothrips hamipes Karny
Diaphorothrips hamipes Karny, 1923 : 296-299. Syntypes 1 2, 2 3 (sic) JAVA: (SMF) [2 2 examined].
The original description refers to one female and two males. However, the only specimens bearing
the original data which have been found are both females although one was labelled as male by
Karny. The species is smaller than both clavipes and unguipes, although the head length varies from
280 to 350 um in the available specimens. Even in the largest specimen the cheek setae are pale
and slender. The external sense cone on the sixth antennal segment is exceptionally large, whereas
it is small in clavipes and only moderately large in unguipes.
SPECIMENS STUDIED.
Jaya: Buitenzorg [Bogor]; 2 2 syntypes under citrus bark, 11.ii.1921 (Kemner) (SMF); 1 ¢ on dead twigs,
18.x.1973 (L. A. Mound) (BMNH). Sumatra: Medan, 1 2? in dry Caesalpinia pods, 15.v.1922 (Fulmek)
(SMF). Malaya: Kuala Lumpur, 3 9, 3 ¢ on dead branches, 26-29.xii.1969 (R. G. & F. Andre) (BMNH).
Diaphorothrips unguipes Karny
Diaphorothrips unguipes Karny, 1920a : 186-189. Syntypes 2 9, SRI LANKA (Ceylon) (SMF) [1 2 examined].
Diaphorothrips thevetii Ananthakrishnan, 1957 : 101-102. Holotype 2, INDIA (TNA) [examined]. Syn. n.
Diaphorothrips spinosus Ananthakrishnan, 1959 : 321-322. Holotype 2, INDIA (TNA) [examined]. [Synony-
mized by Ananthakrishnan & Jagadish, 1970: 275.]
The original description of unguipes refers to one winged and one wingless specimen from Sri
Lanka, but only the winged individual has been studied. D. thevetii was based on a rather small,
damaged female and the original description did not compare this specimen with other members
of the genus. The differences quoted by Ananthakrishnan (19736: 87) are interpreted here as
being related to body size. D. spinosus was based on a male and female with a longer head and
164
Figs 16-21 Males of Dinothrips species. 16, D. spinosus. 17, D. juglandis. 18, D. longicauda.
19-21, antennal segment III, (19) D. spinosus; (20) D. sumatrensis; (21) D. juglandis.
165
with longer setae than the thevetii holotype. However, none of these specimens differs significantly
from the syntype listed below. The date of collection and the collector’s number on the holotype
slide of spinosus differ from those given in the original description.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): Bentotta, 1 2 syntype of unguipes (W. Horn 99) (SMF). India: Madras; 2 holotype
of thevetii, on Thevetia, 19.xi.1955 (Ananthakrishnan); 2 holotype of spinosus from inside dry twigs of
Thevetia, ii1.1957 (Ananthakrishnan); 1 2 on Sesbania twigs, 2.iv.1963 (BMNH); 3 2 on Erythrina twig,
20.1x.1965 (1 2 SMF, 2 2? BMNH).
DINOTHRIPS Bagnall
Dinothrips Bagnall, 19085 : 190. Type-species: Dinothrips sumatrensis Bagnall, by monotypy.
Dinothrips Bagnall; Priesner, 1959 : 52.
Paxillothrips Ananthakrishnan, 1961a: 250. Type-species: Paxillothrips longicauda Ananthakrishnan, by
monotypy. Syn. n.
Dinothrips Bagnall; Ananthakrishnan, 1964 : 88; 19735: 91.
Large, dark, macropterous species of Idolothripini. Head rectangular, about twice as long as broad,
projecting slightly in front of eyes; eyes large and equally developed on dorsal and ventral surfaces; head
with | pair of anteocellar setae, 1 pair of postocular setae and 1 pair of long setae on the vertex (post-
ocular setae pair II); cheeks set with stout spine-like setae. Antennae 8-segmented; segment III about
5-0 to 5-5 times as long as broad, with 2 sense cones, IV with 4 sense cones. Pronotum shorter than head,
almost twice as broad as long, usually with 5 pairs of major setae but sometimes with an additional pair
of posteroangulars (Fig. 24). Fore tarsal tooth present in ¢ and 2, but much larger in 3; fore femora of
3 enlarged with numerous spine-like setae at base (Fig. 16). Mesothoracic anterior angles of 3 with
spiracles produced into a laterally projecting, usually bifurcate, process; this process not developed in
? and some small 3. Pelta triangular, lateral lobes separated (Fig. 30). Tergites II-VII with 2 pairs of
sigmoid wing-retaining setae, those on VII much reduced, tergal accessory setae straight (Fig. 31);
sternite IX of 3 with a pair of large spines (Fig. 33). Tube with straight sides evenly narrowing to apex,
about 4 times as long as broad and about as long as head.
Although Dinothrips is a small genus, only five species now being recognized, the individuals
are frequently large. They are found on dead branches in the area between northern India,
Sri Lanka, New Guinea and the Philippines, and they frequently constitute the dominant element
in the thrips fauna of this habitat. In both Malaya and Java D. sumatrensis and D. spinosus have
been found together on recently fallen tree trunks where they were coexisting in large numbers
with Ecacanthothrips species. Each Dinothrips species is sexually dimorphic, the males having a
pair of enlarged mesothoracic spiracular processes (Figs 22, 26-29) which are not developed in
females. However, the males are subject to allometric growth patterns such that these processes
are large in large individuals but small or absent in small individuals. This sexual dimorphism
and allometry is the cause of the large number of names which have been applied to the two com-
mon species.
The monobasic genus Paxillothrips was distinguished from Dinothrips by the small peg-like
mesothoracic processes, and by the form of the pelta. However, the pelta of the two male and one
female paratypes of /ongicauda which have been studied is similar to that of Dinothrips species
and not like that given in the illustration which accompanied the original description. Moreover,
the mesothoracic process is so variable in Dinothrips, even being absent in some males, that it is
of little practical value as a generic characteristic. Dinothrips is related to both Elaphrothrips and
the monobasic African genus Lamillothrips. In the latter genus the anterior angles of the pronotum
are produced into tubercles in the male, and the lateral lobes of the pelta are not separated. In
Oriental Elaphrothrips species the tergites bear accessory sigmoid setae anterolateral to the major
wing-retaining setae, and the first ocellus is situated anterior to the ocellar setae, on the head
prolongation.
Key to species
1 Antennal segment III with base pale (Fig. 21); mesothoracic spiracular process of 3 small
(Figs 27, 28) . i : ) : ; } ‘ ‘ . ; : : : is
166
— Antennal segment III with base dark (Figs 19, 20); mesothoracic spiracular process of 3
frequently large (Figs 22, 29) : 3
2 Pronotal anteromarginal setae longer than the distance between ‘their bases (Fig. 25); median
pair of setae on vertex small, no longer than cheek setae and about 0:3 times as long as
postocular setae (Fig. 17); cheek setae long and pale (¢ 160-170 um; 2 95-100 um) [India,
Burma] . ; juglandis (p. 167)
— Pronotal anteromarginal setae shorter than distance between their bases; median pair of setae
on vertex about 0-75 times as long as postoculars (Fig. 18); cheek setae short and pale
(50-90 um) [India] . : longicauda (p. 167)
3 Méesothoracic spiracular process oF 3 not bifurcate but with dentate margin (Fig. 29); cheek
setae less than 100 um long [Philippines] . ; : , monodon (p. 167)
— Mesothoracic process of 3 bifurcate or reduced (Figs Des 26) F 4
4 Antennal segment III usually less than 3-5 times as long as the apical dark band on, this segment
(3S 2:2-3-2; 2 2-8-3-9) (Fig. 20); cheek setae long and pale in ¢ (110-180 um) but shorter
and sometimes dark in 2 (60-100 um); anteocellar setae long (¢ 140-180 um; 2 120-160 um);
3 usually, 2 sometimes with more than one pair of major posteroangular pronotal setae
(Fig. 24); epimeral tubercles often well developed; antecostal ridge of sternite II in 2 often
interrupted medially [Malaya to Borneo] . , sumatrensis (p. 170)
— Antennal segment III usually more than 3-5 times as long a as apical dark band on this segment
(3 3-5-4-9; 2 3-4-5-3) (Fig. 19); cheek setae shorter and darker ($ 70-140 um; 2 50-90 um);
anteocellar setae shorter (90-150 um); only one pair of posteroangular pronotal setae (Fig.
16); epimeral tubercles weak even in large 3; antecostal ridge of sternite II in 2 not inter-
rupted medially [India to Solomon Is] . : : : P ; : . Spinosus (p. 169)
Dinothrips juglandis Moulton
(Figs17, 21, 25,28)
Dinothrips juglandis Moulton, 1933 : 6. Holotype 3, INDIA (BMNH) [examined].
This species is readily distinguishable from the others in this genus by having the pronotal
anteromarginal setae longer than the distance between their bases. In the specimens listed below
the cheek setae are long and pale and the mesothoracic spiracular processes are not well developed
(Fig. 28).
SPECIMENS STUDIED.
India: Bengal, Lopchu, holotype 3, paratype 2 (also 3 3 1 2 with identical data) under bark of Juglans
regia, 26.ix.1929 (J. C. M. Gardner) (BMNH). Burma: Mishmi Hills, 2 3, iii.1935 (M. Steel) (BMNH).
Dinothrips longicauda (Ananthakrishnan) comb. n.
(Figs 18, 27, 32)
Paxillothrips longicauda Ananthakrishnan, 1961a: 250. Holotype 9, INDIA (TNA) [not examined].
Two male and one female paratypes of this species have been studied out of an original series
of 4 9, 2 3. These specimens resemble the other species in Dinothrips in most characters, but have
rather paler wings and small peg-like mesothoracic spiracular processes. The available specimens
are small, the head appears to be slightly more concave behind the eyes, and the metanotal
sculpture of the male is developed into a U-shaped ridge (Fig. 32).
SPECIMENS STUDIED.
India: Bombay, Ramangulli, 2 3, 1 2 paratypes in hollow of bamboo, 28.1.1949 (TNA).
Dinothrips monodon Karny
(Fig. 29)
Dinothrips monodon Karny, 1920b : 204. Holotype 3, PHILIpPINEs (SMF) [examined].
The male of this species is recognized easily by the unforked, toothed spiracular process (Fig. 29).
167
Figs 22-29 Males of Dinothrips species. 22-24, D. sumatrensis, (22, 23) mesothoracic spiracular
process. 25, D. juglandis. 26-29, mesothoracic spiracular processes, (26) small D. spinosus; (27)
D. longicauda; (28) D. juglandis; (29) D. monodon.
168
Moulton (1943) records 6 3, 1 2 of monodon from the Philippines, but the original description
was based on a single male.
SPECIMEN STUDIED.
Philippines: Mindanao, Butuan, holotype 3 (Baker) (SMF).
Dinothrips spinosus (Schmutz) comb. n.
(Figs 16, 19, 26, 33)
Ischyrothrips spinosus Schmutz, 1913 : 88. Holotype 2, Skt LANKA (Ceylon) (NM) [examined].
Dinothrips affinis Bagnall, 1915c: 270. LECTOTYPE 3, BorNEo (BMNH), here designated [examined].
Syn. n.
Dicaiothrips crassiceps Bagnall, 1921b : 399. Holotype 2, BURMA (BMNH) [examined]. Syn. n.
Dinothrips jacobsoni Karny, 1921a: 283. Holotype 3, JavA (SMF) [examined]. Syn. n.
Dinothrips kemneri Karny, 1923: 294. LECTOTYPE 3, JAvA (SMF), here designated [examined].
Syn. n.
Dinothrips anodon Karny, 1923: 295. Syntypes 1 3, 1 2, JAvA (depository unknown) [not examined].
Syn. n.
Dinothrips gardneri Moulton, 19285: 290. Holotype 3, INDIA (CAS) [examined]. Syn. n.
Dinothrips malloti Moulton, 1933 : 6. Holotype 3, INDIA (BMNH) [examined]. Syn. n.
Dinothrips celebensis Bagnall, 1934a: 485. Holotype 3, SULAWEsI (Celebes) (MNHN) [examined].
Syn. n.
[Dinothrips sumatrensis Bagnall; Priesner, 1959 : 52. Misidentification.]
D. spinosus is the most commonly collected species in the genus, with an extensive distribution
in tropical rain forest between northern India, Sri Lanka, Sulawesi and New Guinea. However,
throughout much of this range it is often found in company with a second closely related species,
sumatrensis, with which it has previously been confused.
30
aN
YY
cs
AN
S
SN
SNe)
A)
38 gay,
(A
¢
Z
Figs 30-33 Males of Dinothrips species. 30, 31, D. sumatrensis, (30) pelta; (31) tergite IV.
32. D. longicauda, metathorax. 33, D. spinosus, sternite IX.
169
The unique holotype female of spinosus was labelled subsequently by Dr H. Priesner ‘Dino-
thrips sumatrensis Bagn’. This specimen is evidently a member of the genus Dinothrips, but it is
indistinguishable from the female paralectotypes of affinis and, like them, differs from sumatrensis
in having the third antennal segment longer, more slender and paler, in having shorter darker
cheek setae and shorter anteocellar setae, and in having only one pair of posteroangular pronotal
setae. The synonymy of affinis with sumatrensis by Mound (1968) is therefore here refuted.
Bagnall (1915c) states that the Dinothrips collected by G. E. Bryant from dead bark and trees
on Mt Matang in December 1913 comprised both sumatrensis and affinis. He distinguished the
second species by having the third antennal segment ‘more slender’ and ‘distinctly longer than in
sumatrensis . . . rarely darker basally and only narrowly blackish-brown at apex’. The series of
specimens remaining in Bagnall’s collection and bearing the above data comprises 2 9, 5 3
which agree with the description of sumatrensis, and 8 9, 8 g¢ which agree with the description of
affinis. One of these male syntypes is here designated lectotype of affinis.
Bagnall (19216) described crassiceps from a unique female, but this specimen cannot be dis-
tinguished from females of spinosus. The remaining six nominal species listed above in the syno-
nymy of spinosus were all established for different variants of the male mesothoracic spiracular
processes. The holotypes of gardneri and jacobsoni, as well as the lectotype of kemneri designated
here, are merely small individuals with small processes, and the ‘heavy, blunt tips’ of these pro-
cesses in malloti described by Moulton are due to the tips being broken. Moreover, celebensis
lies within the range of variation of spinosus (not of sumatrensis, cf. Mound, 1968), and judging
from the description, this is also true of anodon.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): holotype 2 of spinosus, No. 54, stuck to sap of fallen tree, between wood and bark,
2.i.1902 (Uzel) (NM). India: Malabar, 8 2 (BMNH); Taliparamba, 1 9 (BMNH), 1 9 (SMF); Kumaon,
1 2; Kiruvatti, 4 3, 2 2; Aryankaru, 1 2; Dehra Dun, 1 3, 1 2 (BMNH); holotype 3 of gardneri, Gola-
tappar, 14.xi.1922 (N. C. Chatterjee) (CAS); holotype 3 of malloti, Lachiwala, on log of Mallotus philip-
pensis, 6.i.1929 (BMNH). Burma: holotype 2 of crassiceps, Myawadi, 900 ft [270 m], 24—26.xi.1911
(F. H. Gravely) (BMNH). Vietnam: North, 1 3, 1 2 (BMNH). Malaya: Kuala Lumpur, 13 g, 12 9;
Perak, 1 3, 1 2; Ringlet, 2 ¢; Kuala Trengganu, 1 2 (BMNH); Bentong, 1 ¢; Serdang 1 3, 1 2 (SMF).
Singapore: 4 3, 1 2 (BMNH). Riau (Riouw) Is: 1 2 (SMF). Sumatra: 18 3, 23 9 (BMNH), 17 3, 192
(SMF). Mentawei Is: 3 3, 1 2 (SMF). Java: holotype ¢ of jacobsoni, Semerang, under bark, 1905 (E.
Jacobson) (SMF); lectotype 3 of kemneri, Bogor (Buitenzorg), under Albizzia bark, 10.xii.1920 (N. A.
Kemner) (SMF); Bogor, 24 3, 6 2 (BMNH), 2 3, 3 2 (SMF); Jasinga, 1 3, 2 2 (BMNH); Sindanglaja,
2 3, 1 2 (SMF). Borneo: 1 2 (SMF); Sarawak, 4 3, 5 9; lectotype 3, paralectotypes 7 3, 7 2 of affinis,
Mt Matang 1000 ft [300 m], on dead tree, 16.xii.1913 (G. E. Bryant) (BMNH). Sulawesi (Celebes):
7 3,1 2 (BMNH); holotype ¢ of celebensis (de la Saviniére) (MNHN). Philippines: 1 3, 5 9 (SMF). New
Guinea: 2 2 (BMNH), 1 3 (SMF). Solomon Is: 1 2 (BMNH).
Dinothrips sumatrensis Bagnall
(Figs 20, 22-24, 30, 31)
Dinothrips sumatrensis Bagnall, 1908b : 191. LECTOTYPE 3, SUMATRA (BMNH) [examined].
Dinothrips furcifer Schmutz, 1913 : 36. Holotype 3, SRt LANKA (Ceylon) (lost) [not examined]. [Synony-
mized by Karny 19206: 203.]
Dinothrips fulmeki Priesner, 1959 : 55. Holotype 3, SUMATRA (SMF) [examined]. Syn. n.
The published data for the original material of sumatrensis were ‘Several examples of both sexes
and in all stages, SUMATRA, Amsterdam Museum’, also ‘Three carded males in the Paris
Museum, Benghalis, Sumatra (Maindron 1885)’. The specimens from Amsterdam were stated to
have been sent by J. C. H. Meyere, but recent correspondence with the Zoological Museum,
University of Amsterdam, has established that there are no specimens preserved there from the
type-series. It would have been out of character for Bagnall, in 1908, to return material to a cor-
respondent. However, Karny (19205) states that Bagnall sent to him some specimens of sumatrensis
from the type locality ‘Nias’ — an island off the western coast of Sumatra. This is interesting be-
cause the introduction to Bagnall’s paper (19084: 183) suggests that the type-series was in alcohol,
170
and an old tube has recently been found in Bagnall’s collection labelled by him ‘Nias/K. Jordan’.
Despite the lack of connection between K. Jordan and J. C. H. Meyere this tube may contain the
type-series of sumatrensis. The specimens have now been mounted onto slides and represent two
species, sumatrensis and spinosus. This mixture is probably the cause of subsequent confusion
over the identity of these species. The description of fulmeki by Priesner (1959), for example,
involves a comparison with specimens of ‘sumatrensis’ which were evidently misidentified spino-
sus. The lectotype of sumatrensis designated here is one of the ‘three carded males’, now mounted
onto a slide, from Sumatra, Benghalis. The antennae are lost but the cheek setae are long and pale
and there are 2 pairs of posteroangular pronotal setae.
SPECIMENS STUDIED.
Malaya: 3 3, 10 2? Kuala Lumpur (BMNH); 1 ¢, 9 2 Pahang (SMF); 1 2 Ringlet; 1 3, 1 2 Perak; 1 9,
Kuala Trengganu (BMNH). Riau (Riouw) Is: 2 2 (SMF). Sumatra: lectotype ¢ of sumatrensis Benghalis,
1885 (Maindron) 7004 85; 14 3, 142 (BMNH); holotype 3 of fulmeki Medan, Rimboen, under bark of a
cut-down tree, v.1925 (L. Fulmek), 2 3 paratypes of fulmeki, 2 3, 3 2 with type-data (SMF). Java: '1 3,
1 2? (BMNB). Borneo: Sarawak, 6 3, 3 2 (BMNH), 6 3, 2 2 (SMF).
ELAPHROTHARIPS Bufifa
Elaphrothrips Buffa, 1909 : 162-163. Type-species: Idolothrips coniferarum Pergande, 1896, designated
by Andre, 1940.
Dicaiothrips Buffa, 1909 : 169-170. Type-species: Thrips schotti Heeger, 1852, designated by Bagnall,
1910. [Synonymized by Hood, 1927 : 238-9.]
Elaphrothrips (Elaphoxothrips) Bagnall, 1932 : 516-517. Type-species: K/eothrips athletes Karny, by mono-
typy.
Elaphrothrips (Cradothrips) Ananthakrishnan, 1973a: 273. Type-species: Elaphrothrips (Cradothrips)
insignis Ananthakrishnan, by monotypy.
Macropterous species of Idolothripini; very variable in size, longest individual 4 or 5 times as long as
smallest individual. Head usually elongate, 2:0 to 4-0 times as long as broad, usually with a long projection
in front of eyes; eyes large and equally developed on dorsal and ventral surfaces; head with 1 pair of
long ocellar setae, often as long as the postoculars, situated posterolateral to the first ocellus, 1 pair of
small postocellar setae, 1 pair of postocular setae and 1 pair of long setae on the vertex; some species
have a second smaller pair of setae on the vertex behind the first pair of postocular setae; cheeks set with
stout spine-like setae, particularly in males. Antennae 8-segmented; segment III about 4 to 7 times as
long as broad, with 2 sense cones, IV with 4 sense cones. Pronotum about half as long as head, about
1-5 times as broad as long, with 5 pairs of major setae. Fore femora of 3 enlarged, usually with a stout
sickle-shaped seta at apex; fore tarsal tooth well developed in 3, very reduced or absent in 2. Fore wings
pale or shaded, slightly broadened in the apical half with 25 to 60 duplicated cilia. Pelta broadly tri-
angular or with lateral lobes which are sometimes separate. Tergites II-VII with 2 pairs of sigmoid wing-
retaining setae and several pairs of usually sigmoid accessory setae, but posteromarginal setae of VII
straight; sternites II-VIII with several small accessory setae, sometimes arranged in a row; sternite IX
of males with a pair of large spines (cf. Dinothrips). Tube with straight sides evenly narrowing to apex,
shorter than total head length and not bearing any obvious lateral setae.
Elaphrothrips is a large pantropical genus in which many species show confusing patterns of
structural variation, both between sexes and also between different-sized individuals of the same
sex. At present it is not possible to distinguish between differences due to allometric growth and
differences due to geographical isolation with any great certainty. In the present account only 17
species from the Oriental Region are recognized and 15 new synonyms are established. Several
of these species are widespread, e.g. denticollis, malayensis and spiniceps (Map 1), and detailed
Studies may eventually demonstrate that these are actually species-groups. However, in view of
the intraspecific variation found in some populations, and in view of the few populations which
have ever been sampled and studied satisfactorily, the species concepts adopted here are con-
sidered to be of greater practical value than those adopted in previous studies. Even with the
present analysis it is remarkable how several species have been found to coexist at some sites,
e.g. bakeri, curvipes, denticollis and sensitivus in Malaya and Java (Table 1).
171
Table 1 Elaphrothrips species from (a) Malaya and (b) Java
(a) Malaya
L. A. Mound
Collection 9 39 42 51 81 83 90 104 106 110 111
Nos.
bakeri _ - = ess Sil es | See - - 19 12 ~
curvipes - 23 - - - 13 je. _ Lo 13 13
denticollis 1 - - - Zee: ALG. 8a 7S - - 12,292 ~
jJacobsoni - -
malayensis 6,4 = 192 _ 23,19 = - - - - -
sensitivus -_ -
(b) Java
L. A. Mound
Collection 119 126 162 166 wai 185 192 193
Nos.
bakeri - _ - = = = - 13.3
curvipes - - ~ 1d - -
denticollis = = = 1¢ 19 - - 3S 52
jacobsoni - - - - - -
malayensis = = = fe - -
sensitivus 13 73,69 19 ac 173-1 = 34,69 43,19
Buffa distinguished Elaphrothrips from Idolothrips by the head being slightly produced in front
of the eyes. He removed coniferarum Pergande and flavipes Hood from Idolothrips and included
them in his new genus. He also erected the genus Dicaiothrips for schotti Heeger and the African
species bottegii which have a long head production and a large sickle-shaped seta on the male
fore femur.
In the Austro-Oriental region Elaphrothrips is related to the genus Mecynothrips which has
three pairs of sigmoid wing-retaining setae on tergites III to V; Dinothrips which has straight
tergal accessory setae; Pyrgothrips in which the eyes are prolonged ventrally, and the monotypic
Tiarothrips which has a distinctive head shape with a very long production in front of the eyes.
The first ocellus and anteocellar setae of this latter genus, however, are situated between the
anterior margin of the eyes and not on the prolongation as they are in the other related genera.
The African genus Derothrips is also similar to Elaphrothrips but, as in Pyrgothrips, it has the
eyes prolonged ventrally. At the eastern end of its range (Map 1) Elaphrothrips appears to be
replaced by the closely related genus Mecynothrips.
Elaphrothrips apertus Girault from Queensland, Australia, was transferred to Bolothrips and
synonymized with badius (Hood) by Mound (1974). Elaphrothrips crassiceps Bagnall from Burma
is transferred above to Dinothrips (p. 169) and synonymized with spinosus (Schmutz). Anantha-
krishnan (1973a : 282) synonymized Elaphridia agasthya (Ramakrishna) with crassiceps which he
then designated as the type-species of his genus Elaphridia. However, the unique holotype female
of crassiceps differs from a syntype female of agasthya which has been studied in the shape of the
pelta and the number of pairs of sigmoid wing-retaining setae on the tergites. E. crassiceps has a
pelta with small, separate lateral lobes and tergites with 2 pairs of sigmoid wing-retaining setae.
E. agasthya has an entire pelta without separate lateral lobes and tergites with only 1 pair of
sigmoid wing-retaining setae which are situated on the posterior margin. E. agasthya, the type-
species of Elaphridia, is therefore regarded as a different species from Elaphrothrips crassiceps,
indeed it should be placed in a different tribe. Elaphrothrips thoreauini Girault (1929a) was de-
scribed from an unspecified number of syntypes of unspecified sex from Queensland, Australia.
172
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There is a slide in the Queensland Museum labelled by Girault ‘Elaphrothrips thoreau- | ini
Girault Type 2 / (outer) Taringa, 26th. Jany. 1929 / under bark living gum / trees / Horistothrips |
corticis Girault / Types’. This slide bears 3 9, 1 ¢ under one cover slip and 1 9 under another.
The 2 9, 1 3 grouped together have dark legs with pale knees and tarsi which correspond to the
description of H. corticis. The smallest 2 with these and the 9 under the separate cover slip have
completely pale legs and represent the second species thoreauini. The 2 under the separate cover
slip is here designated as LECTOTYPE and the species should now be referred to as Horistothrips
thoreauini (Girault) comb. n.
The length of the head prolongation is measured laterally from the base of the antenna to the
anterior margin of the eye, and the breadth is measured across the base anterior to the eyes.
Key to the Oriental species
1 Cheek setae shaded, pale to dark . "
— Cheek setae colourless, with no trace of pigment
2 Head prolongation very short, 8 times as broad as long; antennal segment iil less than 5 times
as long as broad; all tibiae dark; basal stem of antennal segments IV and V pale; ° fore
femur with a dark tubercle on inner margin : . _ insignis (p. 181)
Head prolongation 4 or less times as broad as long; 2 fore femur without a tubercle . : 3
3 Antennal segment III 6 times as long as broad; head prolongation 4 times as broad as long;
all tibiae dark; antennal segment III pale with a narrow dark apical ring, basal stem of IV
and V pale. mahensis (p. 182)
— Antennal segment III less than > 2 times as long as ; broad; head prolongation | less than 3-5
times as broad aslong 4
4 Head prolongation 1-8 times as broad as Jong: antennal segment Ul with small area of apical
dark shading, basal stem of IV—VI pale, length of III and IV subequal; apices of tibiae and
tarsi pale, 3 foretarsal tooth slim; pelta (Fig. 49). ; . amoenus (p. 176)
— Head prolongation more than 2 times as broad as long; antennal segment IV-VI with darker
basal stems. 5
5 Hind tibiae with apical half, other tibiae with apices and tarsi pale: antennal segment ill pale
with a slightly shaded apex, segments IV and V with patchy shading in basal stem; major
and minor ¢ with concave inner margin of fore tibiae (Fig. 34) . : :
— All tibiae completely dark; basal stem of antennal segments IV and V evenly dark :
6 Antennal segment III equal to or shorter than IV; distance between base of left ocellar seta
and left postocular seta greater than the length of either of these setae; pelta (Fig. 47)
curvipes (p. 178)
-— Antennal segment III longer than IV; distance between base of left ocellar seta and left
postocular seta less than the length oF either of these setae . : . notabilis (p. 182)
7 Dark apex of antennal segment III extending beyond apical bulge, about one-third length of
segment; head prolongation 2:1 times as broad as long; pelta with lateral lobes separate or
only narrowly joined to centre (Fig. 43); anteroangulars of 3 longer and stouter than small
mid-lateral setae, 2 with posteroangular setae short, little more than 0-5 times total median
length of pronotum (Fig. 41). ‘ jacobsoni (p. 181)
— Dark apex of antennal segment III not extending ‘beyond apical bulge, about one-quarter
length of segment; head prolongation 2-4-3-4 times as broad as long; pelta with lateral
lobes more broadly joined to centre (Fig. 52); anteroangulars of $ shorter than well-
developed mid-laterals, 2 with DON en ees setae long, more than 0-75 times total median
on
sN
length of pronotum . : . bakeri (p. 176)
8 Head prolongation 3-0 to 5: 0 times as broad as ; long: antennal segment III 5-1 or less times as
long as broad; all tibiae completely dark. 9
—- Head prolongation 1-0 to 3-0 times as broad as long; antennal segment ll often longer;
tibiae sometimes partly pale. 11
9 Head prolongation 4-0 to 5-0 times as broad as ; long (Fig. 40); dark apex of antennal segment
III broad, more than one-third length of segment, IV and V sometimes with paler basal
stem; lateral lobes of pelta narrowly joined to centre (Fig. 44) . ; spiniceps (p. 184)
— Head prolongation 3-0 to 4-0 times as broad as long; dark apex of antennal segment III not as ‘
extensive :
10 Antennal segments IV and Vv usually with paler base: lateral lobes of pelta narrowly joined to
centre (Fig. cf. bakeri) ; ; . procer (p. 183)
174
Figs 34-42 Elaphrothrips species. 34, E. curvipes, 3 fore tibia and tarsus. 35, E. denticollis
(mucronatus holotype), 3 fore femur. 36, 37, 3 fore tibiae and tarsi, (36) E. sensitivus holotype 3;
(37) E. denticollis (sumbanus holotype 3). 38, E. denticollis (productus holotype 3). 39, E. greeni
(bouvieri holotype 3). 40, E. spiniceps 9. 41, E. jacobsoni 3. 42, E. bakeri 3, tergite III.
Wee:
- Antennal segments IV and V completely dark; pelta entire (Fig. 48), small species from
Seychelles , . seychellensis (p. 184)
11 All tibiae with apical half pale; antennal segment Il more than 6 times as long as broad,
basal stem of antennal segments IV and V usually slightly paler; head prolongation 1-0 to
1-5 times as broad as long (Fig. 39) . p greeni (p. 180)
— At least mid tibiae completely dark; antennal segment ne usually less than 6 times as long as
broad, if more than 6 times as long as broad, then antennal segments IV and V with dark
basal stem ‘ 12
12 Basal stem of antennal segments ‘IV and Vv pale, yellowish distinctly paler than apex of
segment . : 13
— Basal stem of antennal segments IV and V as dark ; as apex of segment, or light brown, only
slightly paler than apex of segment. 14
13. Basal one-fifth of antennal segment VI clear yellow, distinctly paler than distal part of segment,
antennal segment III distinctly longer than IV; pelta (Fig. 45) —.. . fulmeki (p. 180)
— Antennal segment VI completely dark, antennal segments III and IV subequal in length
malayensis (p. 182)
14 Antennal segment [II uniformly brown . , : ‘ . athletes (p. 176)
— Basal stem of antennal segment III distinctly paler than apex 2 : 15
15 Hind tibiae long, equal to or longer than median head length including the head production;
antennal segment III longer than IV and 4-7 to 6-2 times as long as broad; major pair of
median head setae usually long and stout, four-fifths or more times length of postocular
setae; basal wing setae B, long, B, 1-0 to 1-3 times as long as B,; 3 fore tarsi almost 3 times
as long as broad (Figs 37, 38); pelta (Fig. 46) . : : denticollis (p. 179)
- Hind tibiae short, equal to or shorter than median head length; ‘antennal segment III equal to
or shorter than IV and 4-0 to 5-5 times as long as broad; major pair median head setae
usually short, two-thirds or less times length of postocular setae; basal wing setae B,
short, B, 1:3 to 2:0 times as long as B,; 3 fore tarsi less than 2 times as long as broad
(Fig. 36); pelta (Fig. 50) . : 7 ' : , ; ; ‘ sensitivus (p. 183)
Elaphrothrips amoenus Priesner
(Fig. 49)
Elaphrothrips amoenus Priesner, 1935a: 174. Holotype 3, VIETNAM (NorTH) (SMF) [examined].
This species is known only from the holotype. It has a long head production, dark cheek setae
and pale yellow basal stems of antennal segments II-VI. It is similar only to the stouter fulmeki
which differs in having pale cheek setae.
SPECIMEN STUDIED.
Vietnam (North): Langson, holotype 4, 6.ii.1925 (SMF).
Elaphrothrips athletes (Karny)
Kleothrips athletes Karny, 1923 : 355. Holotype 3, JAvA (SMF) [examined].
Elaphrothrips (Elaphoxothrips) athletes Karny; Bagnall, 1932 : 516-517.
This species is known only from the holotype. It is a major male with a stout fore tarsal claw,
pale cheek setae and a long head production. It is most similar to denticollis but may be dis-
tinguished by the colour of the third antennal segment. The antennae of the athletes holotype
are poorly preserved but segment III appears to be uniformly pale brown.
SPECIMEN STUDIED.
Java: Semarang, holotype 3, ix.1909 (E. Jacobson) (SMF).
Elaphrothrips bakeri (Karny)
(Figs 42, 51, 52)
Dicaiothrips bakeri Karny, 19206 : 206. Holotype 3, PHiLippines (SMF) [examined].
Elaphrothrips mentaweiensis Priesner, 1929b : 201. Holotype 3, MENTAWEI Is (SMF) [examined]. Syn. n.
176
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E. spiniceps; 45, E. fulmeki;
48, E. seychellensis; 49, E. amoenus; 50, E. sensitivus; 51,
depokensis syntype).
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177
,
46, E. denticollis; 47, E. curvipes;
52, E. bakeri (v.
Figs 43-52 Male peltas of Elaphrothrips species. 43, E. jacobsoni;
E. bakeri;
Elaphrothrips bakeri var. depokensis Priesner, 1935a : 159. 10 syntype 3, 2, JAVA (SMF) [1 3, 1 2 examined].
Syn. n.
Elaphrothrips imitator Priesner, 19355 : 249. Holotype 3, JAVA (SMF) [examined]. Syn. n.
This species is dark brown except for antennal segment III which is pale yellow with a small
brown patch at the extreme apex. The head hasarelatively short prolongation in front of the eyes,
and the cheek setae are dark.
The holotype of bakeri is a major male with an enlarged fore femur and a stout fore tarsal
tooth. The holotype of mentaweiensis is a minor male with a slim fore tarsal tooth but is otherwise
inseparable from bakeri.
E. bakeri var. depokensis was described from 10 male and female syntypes differing from
bakeri in the relative lengths of antennal segments IV and V. One major male and one female
with type-data and labelled ‘type’ by Priesner have been examined and are indistinguishable from
bakeri.
The holotype major male and female paratype of imitator which were apparently collected
together have also been examined. They are rather poorly preserved specimens which differ from
bakeri in the diffuse colour of antennal segment III, and in the slightly shorter segment IV and
head prolongation. These characters, however, are not regarded as sufficient to distinguish the
specimens as a separate species.
E. bakeri is most similar to procer. The latter species, from Sri Lanka and India, differs in
having pale cheek setae and pale basal stems of antennal segments IV and V. It may also be
distinguished from the dark brown species jacobsoni by the extent of the apical dark patch on
antennal segment III, the shape of the pelta and the lengths of the pronotal posteroangular setae.
SPECIMENS STUDIED.
Malaya: Negeri, 2 ¢ on Dryobalanops aromatica, 24.vii.1970; Damansara Forest Reserve, 1 2? on dead
branches and leaves, 3.x.1973 (L. A. Mound); Gombak, 2 3 3 on dead leaves on branches, 7.x.1973,
1 2 on dead branch, 13.x.1973 (L. A. Mound); Genting Highlands 3000 ft [1000 m], 2 9, 1 ¢ on dead leaves
14.x.1963, Tea Estate, 3500 ft [1150 m], 1 2, 3 ¢ on dead leaves on branch, 6.x.1973 (L. A. Mound)
(BMNH). Mentawei Is: Siberoet, holotype 3 of mentaweiensis from dried leaves, 24.ix.1924 (H. H.
Karny 101) (SMF); Sipora, 2 9, 2 3 paratypes of mentaweiensis (H. H. Karny 221) (BMNH), | 2 paratype
of mentaweiensis 9.x.1924 (H. H. Karny) (SMF). Java: Jasinga Forest Reserve, 3 2, 1 3 on dead leafy
branch, 30.x.1973 (L. A. Mound) (BMNH); Depok, 1 9, 1 3 syntypes of depokensis on bushes of fresh and
dried leaves, 14.v.1929 (H. H. Karny); Halte Dawoean, holotype 3 with 1 2 paratype of imitator on
Hevea, xi.1906 (De Vos van Nic.) (SMF). Borneo: Sarawak, 1 3 (C. R. Wallace) (BMNH); 6 3, 5 2 on
dead leaves, viii.1972 (Kizasawa & Sohma) (SO, Tokyo). Philippines: Palawan, Puerto Princesa, holotype
3 of bakeri (Baker) (SMF).
Elaphrothrips coreanus Woo
Elaphrothrips coreanus Woo, 1974:69. Holotype °, Korea (Dept. Agric. Biol., Seoul Nat. Univ.,
Suweon) [not examined].
The holotype of this species and the specimen referred to as Elaphrothrips antennalis Bagnall
by Woo (1974) have not been available for study. The descriptions and illustrations, however,
would indicate that neither specimen belongs in this genus.
Elaphrothrips curvipes Priesner
(Figs 34, 47)
Elaphrothrips curvipes Priesner, 1929b : 206. 2 syntype 3, MENTAWEI Is (SMF) [1 ¢ examined].
Elaphrothrips karnyi Priesner, 1935b : 246. Holotype 9, SUMATRA (SMF) [examined]. Syn. n.
Elaphrothrips secus Ananthakrishnan, 1973a: 278. Holotype 2, INDIA (TNA) [examined]. Syn. n.
This species has lightly shaded cheek setae, characteristically patchy shading in the basal stems
of antennal segments IV and V and both major and minor males have a concave inner margin
to the fore tibiae. It is most similar to notabilis which differs in having much longer major head
and pronotal setae.
178
SPECIMENS STUDIED.
India: West Bengal, Kalimpong, holotype 2, paratype ¢ of secus, on dry twigs, 20.iv.1969 (7. N. Anantha-
krishnan) (TNA). Laos: Vang-Viong, 1 3, 1 2 on dead leaves, 21.iii.1975 (S. Yamaguchi) (SO, Tokyo).
Thailand: Ban lung tong, Phuket Is, 2 ¢ on dead leaves, 11.i.1975 (S. Yamaguchi) (SO, Tokyo). Malaya:
Negri Sembilan, 2 9, 1 ¢ on oil palms (Chemara Research Station); Gombak, on dead branches, 3 3, 2 9,
30.ix.-13.x.1973 (L. A. Mound); Genting Highlands 3000 ft [1000 m], 1 ¢ on dead leaves, 14.x.1973
(L. A. Mound); Kuala Lumpur Agricultural University, 1 g on rolled leaves of Pittosporum ferrugineum,
15.x.1973 (L. A. Mound) (BMNH). Sumatra: Fort de Kock, 920 m, 2 holotype of karnyi from a deserted
larval borehole in a plant stem, xi.1920 (E. Jacobson) (SMF). Mentawei Is: Sipora, Sioban, 1 3 syntype
of curvipes from dried leaves, 29.x.1924 (H. H. Karny 221) (SMF). Java: Bogor Gardens, 1 3 on dead
leaves, 25.x.1973, 1 3 on dead creepers, 28.x.1973 (L. A. Mound) (BMNH).
Elaphrothrips denticollis (Bagnall)
(Figs 35, 37, 38, 46)
Dicaiothrips denticollis Bagnall, 1909b : 527. Holotype 2, NiAs (BMNH) [examined].
Elaphrothrips beesoni Ramakrishna, 1934 : 7. Syntypes 3, 9, INDIA (TNA) [2 3 examined]. Syn. n.
Elaphrothrips mucronatus Priesner, 1935a : 167. Holotype 3, JAVA (SMF) [examined]. [Synonymized with
beesoni by Ananthakrishnan, 1973 : 278.]
Elaphrothrips sumbanus Priesner, 1935a: 169. Holotype 3, SuMBA (SMF) [examined]. Syn. n.
Elaphrothrips productus Priesner, 1935a: 170. Holotype 3, SumMBA (SMF) [examined]. Syn. n.
Elaphrothrips productus f. obscuricornis Priesner, 1935a: 171. Syntypes 3, 2, SUMBA (SMF) [1 ¢ ? syntype
examined]. Syn. n.
From recent material collected in Malaya and Java it appears that the males of this species are
very variable. The large males from this material have a slim fore tarsal tooth, a slim fore femur
with a large, pale sickle-shaped seta at the apex and a pair of well-developed median setae on
the vertex. This form corresponds with the holotype male of mucronatus.
A male labelled by Priesner E. productus f. obscuricornis has been examined. It is not labelled
‘type’, but it has the same data as the productus holotype. Since the type-data for f. obscuricornis
was not indicated by Priesner, this specimen is assumed to be a syntype. It has a slim fore tarsal
claw, a slim fore femur with a smaller, darker, sickle-shaped seta at the apex, and a pair of well-
developed median setae on the vertex. This specimen is indistinguishable from the smaller males
in the recently collected series of this species.
The holotype male of sumbanus differs from the previous form in having finer median setae
on the vertex and a longer third antennal segment. This segment is 6-2 times as long as broad in
the holotype, whereas in the other specimens it is only 4-7 to 5-7 times as long as broad.
The holotype male of productus is a large major male with a stout fore tarsal claw, a greatly
enlarged fore femur with a small dark sickle-shaped seta at the apex and fine median setae on
the vertex as in sumbanus. Ananthakrishnan (1973a) regarded productus as a distinct species,
differing mainly in its colouring. The long series of specimens in the British Museum (Natural
History), however, shows this variation in antennal and fore tibial colour.
The holotype of denticollis is a unique female. It is in poor condition with most of the head
setae missing but it has one complete antenna and six legs. It is indistinguishable from the females
associated with the male holotypes of mucronatus, productus and sumbanus, and also from recently
collected females from Malaya and Java.
E. beesoni was described from numerous specimens collected by the Sandal Insect Survey at
Fraserpet, Coorg, 1930. Ramakrishna compares his species with greeni which differs in colour
and the relative lengths of antennal segments III and IV. In greeni they are subequal in length
and in beesoni segment III is longer than IV. The tibiae of beesoni are also dark and differ from
those of greeni, which are distinctly bicoloured. The two male syntypes of beesoni which have
been examined lack antennae but in the shape of the head and fore legs and the colour of the
tibiae they are inseparable from the mucronatus-type males. Ananthakrishnan (1973a) synony-
mized greeni with beesoni but for the above reasons greeni is kept here as a distinct species.
179
However, it is possible that beesoni was based on a mixed series, especially as it was collected
with chandana (see procer).
The characters distinguishing the two males of sumbanus and productus are not regarded as
sufficient to separate these from the remaining material listed below. This is here interpreted as
representing one variable species which is widespread between Sri Lanka and Borneo.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): Peradeniya, 1 3, 25.vii.1913 (A. Rutherford) (BMNH). India: Coorg, 2 3 syntypes of
beesoni on Sandal, 24.x.1930 (TNA). Burma: Mishmi Hills, Delei River 1700 ft [570 m], 1 3, iii.1935
(M. Steele) (BMNH). Malaya: Selangor, 1 9, iii.1898 (Biro); 1 3, on Flindersia brayliana leaves, 3.xii.1956
(Forest Research Institute); Kuala Lumpur, 1 2 on Citrus medica v. acida, 7.ix.1928 (G. H. Corbett);
Gombak, 8 9, 11 3 on dead leaves on branches, 7.x.1973 (L. A. Mound); Genting Highlands 3000 ft
[1000 m], 2 2, 1 3 on dead leaves, 14.x.1973 (L. A. Mound); Genting Tea Estate 3500 ft [1150 m], 29,23
on dead leaves on branch, 6.x.1973 (L. A. Mound) (BMNH). Java: Bogor Gardens, | 3 on dead leaves,
25.x.1973, 1 2 on large dead leaves on tree, 26.x.1973 (L. A. Mound); Jasinga Forest Reserve, 5 2, 3 g on
dead leafy branch, 30.x.1973 (L. A. Mound) (BMNH); Depok, holotype 3 and paratype 2 of mucronatus
on bushes of fresh and dried leaves, 14.v.1929 (H. H. Karny) (SMF). Nias: holotype 2 of denticollis
(K. Jordan) (BMNH). Sumba: Kambera, holotype ¢ and paratype 2 of productus, 1 3 ? syntype of
productus f. obscuricornis (SMF), 3 9, iii.1925, 1 2, 2 3, 6.ii1.1925, 1 3, 12.iii.1925 (Dammerman); Kanang-
gar, 700 m, | 3, v.1925 (Dammerman) (BMNH); Laora 100 m, holotype ¢ and paratype 2 of sumbanus,
v.1925 (Dammerman 89) (SMF). Borneo: 3 3 on dead leaves, viii.1972 (Mizusawa & Soma) (SO, Tokyo).
Elaphrothrips fulmeki Priesner
(Fig. 45)
Elaphrothrips fulmeki Priesner, 1935b : 242. Holotype 3, SUMATRA (SMF) [examined].
This species is known only from the holotype which is a major male. It has a broad fore tarsal
claw, pale cheek setae and pale basal stems to antennal segments III—VI. It is distinguished from
amoenus by the pale cheek setae and segment III being longer than IV. It is most similar to
malayensis and differs only by having a pale base to antennal segment VI which in malayensis is
completely dark. All the males of malayensis that have been examined, however, are small and
it is possible that ful/meki represents the major form of this species. It is also possible that sensitivus
is a larger darker form of malayensis, but these major males differ from fulmeki in having dark
antennal segments IV—VI, a longer more slender fore tarsal claw and a differently shaped pelta.
If malayensis and sensitivus are one species then ful/meki must represent a second species.
SPECIMEN STUDIED.
Holotype 3, Sumatra: Medan (L. Fulmek 24) (SMF).
Elaphrothrips greeni (Bagnall)
(Fig. 39)
Dicaiothrips greeni Bagnall, 1914 : 289. Holotype 3, Skt LANKA (Ceylon) (BMNH) [examined].
Dicaiothrips bouvieri Vuillet, 1914: 276. Holotype 3, INDIA (BMNH) [examined]. [Synonymized by
Mound, 1968 : 96.]
Elaphrothrips micidus Ananthakrishnan, 1973a: 275. Holotype 2, INDIA (TNA) [not examined]. Syn. n.
The holotype of greeni lacks antennae and has only one fore leg which is without tarsi. Abdominal
segment VII is missing and the detached segments VIII—X are from a female. The fore femur,
however, is enlarged as in males and it would appear that the detached abdominal segments are
from another specimen. One unidentified male in the British Museum (Natural History), mounted
and labelled by Bagnall ‘CEYLON / E. E. Green 2821’ appears to be the same species. This,
also, is without antennae but does have two complete fore and hind legs and one complete mid
leg. The tarsi and apical half of all tibiae are pale. The original description would suggest that
antennal segments IV and V have a slightly paler stem. Three other specimens in the British
Museum (Natural History), two males and one female from India, have all tibiae and antennal
180
segments III-VI distinctly bicoloured. The holotype of bouvieri has bicoloured tibiae but the basal
stems of antennal segments IV—VI are dark. The 3 paratype of bouvieri has darker fore tibiae
and tarsi but antennal segments IV and V have patchy, somewhat paler shading on the basal
stem. Mound (1968), recognizing this variability in colour, synonymized bouvieri with greeni.
The male and female paratypes of micidus which have been examined have all tibiae bicoloured
and pale basal stems to antennal segments III-V. They are therefore here synonymized with
greeni. The relationship of this species to beesoni (cf. Ananthakrishnan, 1973a) is discussed under
denticollis.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): Peradeniya, holotype 3 of greeni on decayed Phaseolus pods (E. E. Green 3023);
1 3, ‘CEYLON’ (E. E. Green 2821), no further data (BMNH). India: Nilgiri Hills, holotype and paratype
3 of bouvieri, Coonoor 2000 m, 14-31.vii.1901 (M. Maindron), 1 3, Mysore (Campbell); Kiruvatti, 1 2
on dry leaves and twigs, 17.ix.1969 (7. N. Ananthakrishnan); Trichur, 1 3 on Areca sheath, 17.viii.1966
(T. N. Ananthakrishnan) (BMNH); Yelagiri, 1 3, 1 2 paratypes of micidus on dry leaves, 8.x.1970
(T. N. Ananthakrishnan) (TNA).
Elaphrothrips insignis Ananthakrishnan
Elaphrothrips (Cradothrips) insignis Ananthakrishnan, 1973a: 273. Holotype 2, INDIA (TNA) [not ex-
amined].
Ananthakrishnan erected a subgenus for this new species described for two females and two
males. It is characterized by the presence of a tooth on the inner margin of the female fore femur.
The head prolongation is very short, about 8 times as broad as long, antennal segment III is less
than 5 times as long as broad and longer than IV, antennal segments III-V have pale basal stems
and all tibiae are dark brown. The two specimens examined are assumed to be paratypes although
the published date of collection was 19.xi.1970.
SPECIMENS STUDIED.
India: Almora, U.P., 1 3, 1 2 paratypes on dry twigs, 19.x.1970 (TNA).
Elaphrothrips insularis Priesner
Elaphrothrips insularis Priesner, 1928a: 57. Holotype 3, JAvA (Hamburg Museum, destroyed).
This species was described from a unique male which was destroyed in the Hamburg Museum.
It was not illustrated and the original description is insufficient to distinguish it from, or synony-
mize it with, other species in the genus from the Oriental Region. However, it would appear to
resemble most closely fulmeki with the head prolongation 1-9 times as broad as long, antennal
segments III-VI with pale basal stems, segment III longer than IV, and a stout fore tarsal claw.
The name insularis is best regarded as a nomen dubium at present.
Elaphrothrips jacobsoni Priesner
(Figs 41, 43)
Elaphrothrips jacobsoni Priesner, 1935b : 243. Holotype 3, SUMATRA (SMF) [examined].
This is a very large black species, similar to bakeri, with dark cheek setae. It may be distinguished
mainly by the more extensive dark apex of antennal segment III, the slightly shorter head pro-
longation and the lateral lobes of the pelta being separate or only narrowly joined to the centre.
SPECIMENS STUDIED.
Malaya: Ringlet, 1 ¢ on dead branch, 12.x.1973; Gombak, 1 2 on dead branch, 13.x.1973; Genting
Highlands, 3000 ft [1000 m], 1 ¢ on dead leaves, 14.x.1973 (L. A. Mound) (BMNH). Sumatra: Fort de
Kock, 920 m, holotype 3 and 1 2 paratype on plant detritus, vii.1922 (E. Jacobson) (SMF). Borneo: | 3
under bark, 1973 (7. Kubayashi) (SO, Tokyo).
181
Elaphrothrips mahensis (Bagnall)
Dicaiothrips mahensis Bagnall, 1921a: 283. Holotype 3, SEYCHELLES (BMNH) [examined].
Dicaiothrips rex Bagnall, 1921a: 281. Holotype 3, SEYCHELLES (BMNH) [examined]. [Synonymized by
Mound, 1968 : 97.]
Dicaiothrips hystrix Bagnall, 1921a: 284. Holotype 2, SEYCHELLES (BMNH) [examined]. [Synonymized
by Mound, 1968 : 97.]
This species has a short head prolongation, a long antennal segment III and a distinctive antennal
colour. Segment III is very pale with a thin dark apical ring and segments IV and V have pale
basal stems with a thin dark basal ring. The holotypes of mahensis and rex represent the major
male form and Aystrix represents the female form of this species. It is most similar to procer which
differs in having pale cheek setae and not so distinctly bicoloured antennal segments IV and V.
SPECIMENS STUDIED.
Seychelles: Mahé, nr Morne Blanc, holotype 3, 1 $ paratype of mahensis, ix—x.1908 (H. Scott); Cascade
hystrix, holotype 3 of rex [6.ix.1908 (H. Scott)], 1 3 paratype of hystrix, 1908 (Seychelles Expedition),
Mare-aux-Cochons, | 9, ix.1908 (H. Scott) (BMNH).
Elaphrothrips malayensis (Bagnall)
Dicaiothrips malayensis Bagnall, 1909) : 525. Holotype 3, Nias (BMNH) [examined].
Dicaiothrips bruneitarsis Schmutz, 1913 : 80. Holotype 3, Ski LANKA (Ceylon) (NM) [examined]. Syn. n.
Dicaiothrips bruneitarsis var. levis Schmutz, 1913: 82. ? Holotype 2, SRr LANKA (Ceylon) (NM) [ex-
amined]. Syn. n.
Elaphrothrips coronatus Bagnall, 1934b: 631. Holotype 2, Sr1 LANKA (Ceylon) (BMNH) [examined].
Syn. n.
The 2 holotypes of bruneitarsis var. levis and coronatus from Sri Lanka are indistinguishable from
a series of females collected in Malaya. These females were collected with males indistinguishable
from the holotype ¢ of malayensis from the Isle of Nias, Java, and the holotype 3 of bruneitarsis
from Sri Lanka. Both sexes have antennal segment III and the basal stems of IV and V pale
yellow, and the legs are dark except for the tarsi. The females have a small fore tarsal tooth and
a slightly enlarged fore femur with a rugose inner margin. These three species are, therefore,
here synonymized.
E. malayensis is most similar to the larger darker species sensitivus in which the basal stems of
antennal segments IV and V are as dark as, or only slightly paler than, the apex of those segments.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): Peradeniya, holotype 3 of bruneitarsis, Nr 64 from the upper side of a banana leaf
in the Botanic Gardens of Peradeniya, 4.i.1902 (Uzel); ? holotype 2 of bruneitarsis var. levis, Nr 83>
10.i.1902 (Uzel) (NM); holotype 2 of coronatus, on banana 2.vi.1913 (A. Rutherford); 2 2, 1g on dead
Hevea leaves, 6.v.1914 (A. Rutherford) (BMNH); Deniyaya, 1 3, 1 2 on dead leaves, 7.xi.1974 (S. Yama-
guchi) (SO, Tokyo). Laos: Van-Viong, 2 3, 1 2 on dead leaves, 21.iii.1975 (S. Yamaguchi) (SO, Tokyo).
Thailand: East, Chanta buri, 1 3, 1 2 on dead leaves, 30.iii.1975; South, Ban lung tong, Phuket Is, 1 3
on dead leaves, 11.i1.1975 (S. Yamaguchi) (SO, Tokyo). Malaya: Negri Sembilan, 1 2 on oil palms (Chamara
Research Station); 11 miles (17 km] Genting Highlands, 1 2, 1 3 on dead leaves, 27.ix.1973 (L. A. Mound);
5000 ft [1700 m] Genting Highlands, 1 @ on dead branch, 1.x.1973 (L. A. Mound); 3500 ft [1200 m]
Genting Tea Estate, 1 9, 2 ¢ on dead leaves on branch, 6.x.1973 (L. A. Mound). Nias: holotype and 2
paratype ¢ of malayensis (K. Jordan) (BMNBH).
Elaphrothrips notabilis Ananthakrishnan
Elaphrothrips notabilis Ananthakrishnan, 1973a : 276. Holotype 2, INDIA (TNA) [examined].
This species has pale shaded cheek setae and the fore tibiae of the males have a concave inner
margin. It is very similar to curvipes but can be distinguished from the latter species by its
unusually long major head and pronotal setae.
182
SPECIMENS STUDIED.
India: Mysore, Kiruvatti Forest, holotype 2, paratype 3 on decaying bark, 6.ix.1967 (JT. N. Ananthakrish-
nan) (TNA).
Elaphrothrips procer (Schmutz)
Dicaiothrips procer Schmutz, 1913 : 73. Holotype 3, Skt LANKA (Ceylon) (NM) [examined].
Dicaiothrips novus Schmutz, 1913 : 76. Holotype 2, Sk1 LANKA (Ceylon) (NM) [examined]. [Synonymized
by Ananthakrishnan, 1973a: 281.]
Dicaiothrips dallatorensis Schmutz, 1913: 77. Holotype 3, Sr1 LANKA (Ceylon) (NM) [examined].
[Synonymized by Ananthakrishnan, 1973a: 281.]
Dicaiothrips proximus Bagnall, 1914: 289. Holotype 3, Sri LANKA (Ceylon) (BMNH) [examined].
[Synonymized by Mound, 1968 : 98.]
Elaphrothrips achaetus Bagnall, 19346: 633. Holotype 2, Srr LANKA (Ceylon) (BMNH) [examined].
[Synonymized by Mound, 1968 : 98.]
Elaphrothrips approximatus Bagnall, 19346 : 635. Holotype 3, Srt LANKA (Ceylon) (BMNH) [examined].
[Synonymized by Mound, 1968 : 98.]
Elaphrothrips chandana Ramakrishna, 1934: 9. Holotype 3, INDIA (unknown) [not examined]. [Synony-
mized by Ananthakrishnan, 1973a: 28.]
Elaphrothrips eranthemi Seshadri & Ananthakrishnan, 1954 : 224. Holotype 3, INDIA (TNA) [1 3 paratype
examined]. [Synonymized by Ananthakrishnan, 1973a: 281.]
This species has a relatively short head prolongation in front of the eyes, paler basal stems to
antennal segments IV and V and distinct lateral lobes to the pelta.
The major males of this species have an enlarged fore femur and a stout fore tarsal tooth. The
minor males have a slim fore femur and a slender fore tarsal tooth. Mound (1968) synonymized
the major male proximus, the minor male approximatus and the female achaetus with procer
which was based on a major male. The holotype female of novus, the holotype minor male of
dallatorensis and a paratype major male of eranthemi are indistinguishable from achaetus,
approximatus and proximus respectively. They are here, therefore, all regarded as synonyms of
procer. E. chandana was described from a few males and females collected with beesoni. They are
smaller and darker and, according to Ananthakrishnan, are synonymous with the ‘dallatorensis’
form.
This species from Sri Lanka and India is similar to bakeri from Malaya apart from the pale
cheek setae and paler basal stems to antennal segments IV and V. The female and minor male
from Laos, however, have almost uniformly dark basal stems to segments IV and V.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): holotype 3 of procer, holotype 2 of novus, Nr. 34, on a bush 19.xii.1901 (Uzel) (NM);
Peradeniya, holotype 3 of dallatorensis, Nr. 83, 10.i.1902 (Uzel) (NM), holotype 3 of proximus, holotype
2 of achaetus, 1 2, 1 3, on pods of Crotalaria, xi.1912 (E. E. Green 3180), holotype 3 of approximatus on
dead Hevea leaves, 6.v.1914 (A. Rutherford 321), 1 3, 21.vii.1913 (A. Rutherford), 1 2 without data
(BMNH). India: Madras, 1 paratype 3 of eranthemi, Teynampet Botanical Gardens, on Eranthemum,
17.1.1953 (T. N. Ananthakrishnan 150) (TNA), 1 2 on Cassia twig, 23.x.1966, 1 2, 1 ¢ on dry twigs,
4.i11.1967, 3 9, 3 g on Sesbania twigs 3.iv.1963; Adichanallur, 1 2 on Areca sheath, 29.viii.1966 (7. N.
Ananthakrishnan) (BMNH). Laos: Vang-Viong, 1 g, 1 2 on dead leaves, 21.iii.1975 (S. Yamaguchi)
(SO, Tokyo).
Elaphrothrips sensitivus Priesner
(Figs 36, 50)
Elaphrothrips sensitivus Priesner, 1929b : 204. Holotype 3, MENTAWEI Is (SMF) [examined].
The holotype of sensitivus is a major male. This species has pale cheek setae and a long head pro-
duction, and the males have a well-developed but slim fore tarsal tooth. It is very similar to
malayensis which may be distinguished by the pale basal stems of antennal segments IV and V.
183
SPECIMENS STUDIED.
Mentawei Is: Sipora, Sioban, holotype 3, from dead leaves, 9.x.1924 (H. H. Karny 164) (SMF). Java:
Bogor, | 2 (labelled by Priesner, allotype of sensitivus, but not referred to in the original description) on
dried leaves, 22.ix.1922 (Soerijat) (SMF); Bogor Gardens, | ¢ on dead leaves of Brownea sp. tree,
18.x.1973, 6 2, 7 $ on dead leaves on tree 19.x.1973, 1 9, 24.x.1973, 4 ¢ on dead leaves, 25.x.1973, 12 9,
18 ¢ on large dead leaves on tree, 26.x.1973, 6 2, 3 ¢ on dead leafy branch, 29.x.1973 (L. A. Mound);
Jasinga Forest Reserve, 1 2, 4 3 on dead leafy branch, 30.x.1973 (L. A. Mound) (BMNH).
Elaphrothrips seychellensis (Bagnall)
(Fig. 48)
Dicaiothrips seychellensis Bagnall, 1921a: 280. Holotype 9, SEYCHELLES (BMNH) [examined].
This is a small dark species from the Seychelles with pale cheek setae. The males have a well-
developed fore femur with a large sickle-shaped seta and a well-developed fore tarsal tooth.
SPECIMENS STUDIED.
Seychelles: holotype 9, Mahé [above Port Glaud, 500-1000 ft [200-300 m], 5.xi.1908 (H. Scott)], 1 3
paratype (Seychelles Expedition 1908-1909), above Port Glaud 500-1000 ft [200-300 m], 1 2 paratype,
5.xi.1908 (H. Scott), Cascade Estate, 800-1000 ft [270-300 m], 1 9, 1.1909 (H. Scott); Silhouette nr Mont
Pot-a-Eau, 1500 ft [500 m], 1 3, viii.1908 (H. Scott) (BMNH).
Elaphrothrips spiniceps Bagnall
(Figs 40, 44)
Elaphrothrips spiniceps Bagnall, 1932 : 514. Holotype 2, INDIA (BMNH) [examined].
Elaphrothrips graveleyi Bagnall, 19346 : 628. Holotype 9, INDIA (BMNH) [examined]. Syn. n.
Elaphrothrips clarispinis Priesner, 19355 : 247. Holotype 3, JAVA (SMF) [examined]. Syn. n.
This is a large, stout, black species with a broad head, only about twice as long as broad, pale
cheek setae and a short prolongation in front of the eyes. The small lateral lobes of the pelta are
narrowly joined to the centre. The holotype of spiniceps, although described and labelled as a
female, is, in fact, a male with enlarged fore femora and a well-developed fore tarsal tooth.
Both antennae are unfortunately missing distal to the second segment.
The holotype of graveleyi is a female with the head and pelta shape as in spiniceps. The basal
stem of antennal segment III is clear pale yellow and segments IV and V have a slightly paler
basal stem. An additional male and female in the British Museum (Natural History) collection
from Burma and two males and two females from Nepal (SO, Tokyo) all have pale basal stems
to segments III-V. The sickle-shaped seta on the fore femur of even the largest of these males is
small.
The male holotype of clarispinis is indistinguishable from the holotype of spiniceps in the shape
of the head and pelta. Both antennae of clarispinis are complete, however, and the basal stem of
segment III has patchy dark shading and the basal stem of IV is hardly paler than the apex.
The fore femur also bears a well-developed sickle-shaped seta. Four males and one female from
Taiwan also have this antennal colouration, and a large sickle-shaped seta in the males, but this
is considered to be no more than regional variation and all these specimens are here regarded as
spiniceps. Ananthakrishnan (1973a) records two males and two females of clarispinis from India.
However, from the redescription and illustrations it is apparent that these specimens have been
misidentified and most closely resemble denticollis which is found in India.
SPECIMENS STUDIED.
India: Sikkim, Gangtok, holotype 2 of spiniceps (Lt. Col. F. M. Bailey), holotype ° of graveleyi, Kurs[eJong
4700 ft [1800 m] with Neoheegeria fumipennis Bagnall, 26.iii.1910 (F. H. Graveley) (BMNH). Nepal:
Ghasa, 2 3, 2 2 on dead leaves, 4.vi.1974 (S. Yamaguchi) (SO, Tokyo). Burma: Mishmi Hills, 1 3, Delei
River, 4700 ft [1800 m] iii.1935; 1 g, Monodon (M. Steele) (BMNH). Taiwan: Nanshanchi, 4 3, 1 2 on
dead leaves, 1.vi.1972 (S. Okajima) (SO, Tokyo). Java: Kamodjan, above Garoet, 1200 m, holotype ¢
of clarispinis, 30.v.1923 (Siebers) (SMF).
184
j i jUNci. ; 55, 56, antennal
Figs 53-58 Machatothrips species. (53) M. biuncinatus; (54) M. antennatus. oo.
sk a0 III and IV, (55) M. quadrudentatus; (56) M. biuncinatus. (57) M. braueri; (58) M. heveae.
185
MACHATOTHRIPS Bagnall
Machatothrips Bagnall, 19085 : 189. Type-species: Machatothrips biuncinatus Bagnall, by monotypy.
Adiaphorothrips Bagnall, 1909b : 536. Type-species: Adiaphorothrips simplex Bagnall, by original desig-
nation. [Synonymized by Priesner 1939: 75.]
Cnestrothrips Priesner, 1932 : 344 (1939 : 75). Type-species : Cnestrothrips dammermani Priesner, by original
designation. Syn. n.
Large, dark, macropterous species of Cryptothripini. Head rectangular, about 1-5 times as long as broad,
projecting only slightly in front of eyes; eyes equally developed on dorsal and ventral surfaces; head with
1 pair of elongate ocellar setae situated posterolaterally to anterior ocellus and outside ocellar triangle,
also 1 pair of postocular setae and 1 median pair of setae on vertex (postocular pair II); cheeks with stout
spine-like setae. Antennae 8-segmented, segment III 2:5 to 5-0 times as long as broad with 2 sense cones,
IV with 4 sense cones. Pronotum shorter than head, twice as broad as long with 5 pairs of major setae.
Fore femora of males and females enlarged, females always and males rarely with a row of teeth on inner
margin; fore tarsal tooth present in ¢ and 9, larger in g. Wings broad, slightly broader in apical half,
pale or dusky, with numerous duplicated cilia. Pelta broadly triangular. Tergites II—-VII with 1 pair of
wing-retaining setae; sternites II-VIII with a row of small accessory setae; sternite LX of males with a pair
of long stout setae. Tube with straight sides evenly narrowing to apex, 3-0 to 3:5 times as long as broad,
about as long as head.
Twelve species are here recognized in this genus and a further two nominal species were not
available for study, but the patterns of inter- and intraspecific variation are difficult to interpret
(Table 2). The species are distinguished on the lengths of the setae on the head and pronotum,
the length and colour of the third antennal segment, and the form and number of tubercles on
the fore femora of the females. Unfortunately none of these characters appears to be stable, and
since it is not unusual for two or more species to be collected together the possibility of hybridi-
zation must be considered in future studies in the field. The species are not infrequently collected
but apparently do not usually exist in large populations. This widespread dispersion in low
numbers suggests a high vagility, and this biological characteristic would contribute to taxonomic
problems. Certainly the taxonomic conclusions which have been reached in this genus, for example
only two new synonyms are established, are in marked contrast to the conclusions reached in
Elaphrothrips and Dinothrips. In both of these genera it is not unusual for each species to form
large populations and, despite the problems associated with allometric growth, there is less over-
lap in the ranges of structural variation between species.
Machatothrips species are widespread between West Africa and Guam throughout the Old
World tropics. The genus is unusual in that it is the females, not males, which bear the obvious
sexually dimorphic characters on which species are defined. However, in both of the new species
described below, decorus and /Jentus, one or more males are known which have female-like fore
legs.
Adiaphorothrips pallicornis Karny (1924) from Queensland is not a member of Machatothrips.
The holotype has slender maxillary stylets and is a member of the Phlaeothripinae. This species
is here transferred to the genus Akainothrips Mound (1971a) and should be referred to as Akaino-
thrips pallicornis (Karny) comb. n. The holotype female of Machatothrips isshikii Ishida (1932 : 7-9)
(HUS) has been examined. The specimen has been remounted and relabelled by Dr Haga as
Acanthothrips nodicornis (Reuter) although this synonymy does not appear to have been pub-
lished.
The monotypic genus Cnestrothrips was erected by Priesner for his species dammermani which
is here shown to be a synonym of Machatothrips antennatus. It differs from the other species of
Machatothrips in the presence of a group of spines on the fore angles of the pronotum, and also
in having a series of small tubercles, without any large teeth, on the inner margin of the female
fore femora. The arrangement of these teeth in M. decorus is intermediate between that of
antennatus and the other species in the genus in having one larger basal tooth and a row of 12 to
14 almost uniform small teeth.
Machatothrips is closely related to Macrothrips from New Guinea although the females of the
latter do not have a row of teeth on the fore femora. Both males and females of Macrothrips
have a hook on the ventral surface of the fore angles of the pronotum. Larger specimens of this
186
Table 2 Variation in the length of antennal segment III and the postocular setae of Machatothrips
species
Antennal segment III Postocular setae
Species length/breadth pair I length/pair II length Distribution
sylvaticus 2°4 to 2:7 3-4 to 5-0 India
quadrudentatus 2°5 10-0 New Guinea
antennatus 2°8 to 3:6 7:0 to 9:3 Malaya
indicus 3-0 48 India
haplodon 3-2 to 3°6 5-0 to 10-0 Africa
lentus 3-2 to 3-8 3-0 to 6°5 Malaya
decorus 3-2 to 3:8 2°8 to 3:3 Malaya
heveae 3-5 to 4:0 2:0 to 2:2 Malaya
artocarpi 3°5 to 4-4 1-9'to 3-7 Japan to Solomon Is
braueri 3-0 to 4:5 I-9to 3°2 Africa
celosiae 4:7 2°4 Taiwan
biuncinatus 4-1 to 5:0 1-4 to 2:8 Malaya to New Guinea
genus have a nodular swelling on the inner margin of the fore femora near the base, they also
have large coxal and epimeral tubercles, and the fore tibiae have from one to three teeth on the
inner margin near the apex. The pronotum of Machatothrips lacks the hook and tubercles, and
only in males of antennatus is a small tooth visible at the apex of the fore tibiae. The tube of
Macrothrips is longer than in Machatothrips being more than four times as long as broad and the
pores on the pelta are situated further forward (Figs 71-78). Partial revisions of Machatothrips
have been published by Mound (1968; 1970).
Key to species
1 Postocular setae pair I 5 to 10 times as long as postocular setae pair 14 ae F : , g
— Postocular setae pair I 1 to 5 times as long as postocular setae pair II : >
2 Antennal segment III yellow, 2:5 times as long as broad (Fig. 55); anteromarginal setae equal
in length to anteroangulars [New Guinea] . : quadrudentatus te 195)
— Antennal segment III as dark as IV, 2°8 to 3-8 times as long as broad F 3
3 Fore angles of pronotum with a group of spine-like setae (Fig. 54); fore femora of 2 with a
complete row of small tubercles (Fig. 66); anteromarginal setae 0-4 to 0-7 times as long as
anteroangulars [Malaya, Borneo] : : antennatus (p. 189)
— Fore angles of pronotum without spine-like setae; fore femora of 2 with a row of 4 to 8 stout
teeth; anteromarginal setae 0:6 to 1:1 times as long as anteroangulars_ . : 4
4 Fore tibiae of 2 with a distinct hump, fore tarsal claw small, fore femora with teeth restricted
to distal half (Fig. 61) [Malaya] ; . lentus (part) (p. 194)
— Fore tibiae of 2 without a hump, fore tarsal claw well developed and sharply pointed, fore
femora with row of teeth extending into basal half (Fig. 60) [Africa]. haplodon (p. 193)
5 Antennal segment III 2-4 to 3-0 times as long as broad at apex ; : ‘ : 6
— Antennal segment III more than 3-0 times as long as broad. . . ,
6 Antennal segment III slightly paler than IV; fore tibiae of 2 without ‘hump [India]
silvaticus (p. 195)
— Basal stem of antennal segments III—VI slightly paler than aoe half; fore tibiae of 2 with a
distinct hump bearing spinules (Fig. 59) [India] . : . indicus (p. 194)
7 Pronotal anteromarginal setae short, less than 0:5 times as long : as anteroangulars: postocular
setae pair II long, pair I 1-4 to 2:8 times as long as pair II (Fig. 53); antennal segment III
long, 4:1 to 5-0 times as long as broad (Fig. 56); distal teeth on fore femora of 2 usually
fused into a ridge (2+3 to 4+5) (cf. Fig. 61) [Malaya to New Guinea] . biuncinatus (p. 190)
— Pronotal anteromarginal setae longer, more than 0:5 times as long as anteroangulars;
antennal segment III shorter, usually less than 4-5 times as long as broad; teeth on 2 fore
femora usually separate . 8
8 Pronotal anteromarginal setae longer than anteroangulars; antennal segment Tl 4-7 times as
long as broad [Taiwan] . ; : ; . . ‘ : ; . celosiae (p. 192)
187
Figs 59-70 Machatothrips species. 59, M. indicus °, fore tibia and tarsus. 60, M. haplodon 9. 61,
M. lentus 2. 62-66, 2 fore femora, (62) M. artocarpi; (63) M. heveae; (64) M. braueri; (65) M.
decorus ; (66) M. antennatus. 67-70, 3 fore tarsi, (67) M. artocarpi; (68) M. decorus; (69,70) M.
lentus (69 gynaecoid 3).
188
- Pronotal anteromarginal setae 0-5 to 1-0 times as long as anteroangulars; antennal segment
III 4-5 or less times as long as broad . é 7
9 Postocular setae pair I 3-0 to 6:5 times as long as pair ti antennal segment tl 3° 2 to 3- 8 times
as long as broad; fore tibiae of 2 with a distinct hump, arrangement of teeth on fore femora
similar to biuncinatus (2+3 to 4+ 6) (Fig. 61) [Malaya] : . lentus (part) (p. 194)
— Postocular setae pair I usually less than 3-2 times as long as pair II, if more than 3-2 times as
long as pair II then antennal segment III 3-5 or more times as long as broad; fore tibiae of
2 without a distinct hump. ' 10
10 Pronotal posteroangular setae short, median pronotal length 1: 8 to 3: 0 times as long as s these
setae (Fig. 58); antennal segment III 3-5 to 4-0 times as long as broad; fore femora of 2
with 5 to 10 teeth throughout its length, first tooth in basal half smaller than second
(Fig. 63) [Malaya, Java] . : : . heveae (p. 194)
— Pronotal posteroangular setae longer (Fig. 57), median pronotal length usually 1-1 to 2:0
times as long as these setae, if more than 2-0 then antennal segment III only 3-2 times as
long as broad . ; 11
11 Tube short, 2-8 to 3-3 times a as long as basal breadth: fore femora of 9 with 5 to 10 closely set
teeth evenly See in size, rarely with a smaller tooth or tubercle in basal half (Fig. 64)
[Africa] . : ; i , . braueri (p. 190)
— Tube longer, 3-3 or more times as Jong ¢ as basal breadth . : 34
12 Fore femora of 2 with a distinctive row of 12 to 14 teeth in distal half (Fig. 65); pronotal
anteromarginal setae 0-5 times as long as ue aaa antennal segment III 3-2 to 3-8
times as long as broad [Malaya] : . decorus (p. 193)
— Fore femora of 2 with 3 to 5 separate teeth in . distal half (Fig. 62); pronotal anteromarginal
setae 0-6 to 0-9 times as long as anteroangulars; antennal segment III 3-5 to 4-4 times as
long as broad [Japan to Solomon Is] . : é 2 : ‘ : artocarpi (p. 189)
Machatothrips antennatus (Bagnall)
(Figs 54, 66, 74)
Adiaphorothrips antennatus Bagnall, 1915b : 594. Lectotype 2, BoRNEO (BMNH) [examined].
Cnestrothrips dammermani Priesner, 1932: 344 (1939: 76). Holotype °, RIAU (Riouw) Is (SMF) [ex-
amined]. Syn. n.
This is a distinctive species, both males and females having a group of spine-like setae on the fore
angles of the pronotum and short postocular setae pair II. The females have a complete row of
12 to 20 small tubercles on the fore femora, and the tergal accessory setae of both g and 9 are
rather robust. The holotype of dammermani has these characteristics and is indistinguishable
from antennatus with which it is here synonymized. This species is most similar to decorus which
may be distinguished by having much longer postocular setae pair II.
SPECIMENS STUDIED.
Malaya: Gombak, dead branch, 2 9, 1 3, 30.ix.1973; 1 3, 7.x.1973; Damansara Forest Reserve, 8 miles
[13 km] north of Kuala Lumpur, 1 9, 3.x.1973 (L. A. Mound) (BMNH). Singapore: Bukit Timah, 2 9,
1 3 on dead branches, 3.xi.1973 (L. A. Mound); 2 3 on Dialium wallichii fallen tree, 16.xi.1973 (D. H.
Murphy) (BMNH). Riau (Riouw) Is: Doerian, holotype 2 of dammermani, caught in low herbage,
vi.1923 (Dammerman 33) (SMF). Borneo: West Sarawak, lectotype 2 of antennatus, Mt Matang, 1000 ft
[300 m], under bark of dead tree, 7.xii.1913; Quop, 1 3, 28.iii.1914 (G. E. Bryant) (BMNH).
Machatothrips artocarpi Moulton
(Figs 62, 67, 71)
Machatothrips artocarpi Moulton, 1928a : 322. Holotype 2, TAIWAN (CAS) [examined].
This dark but otherwise unremarkable species is recorded from Japan, Formosa, Guam, Philip-
pines, New Guinea and the Solomon Is, but not, as yet, from further west.
SPECIMENS STUDIED.
Japan: Ishigaki-jima I., Mt Banna, 2 2, 2 ¢ on dead twigs, 16.vi.1972 (S. Okajima) (SO, Tokyo). Taiwan
(Formosa): Kagi, holotype 2, paratype 3, on Artocarpus, 10.viii.1927 (R. Takahashi) (CAS). Guam: | 3
189
under dead bark (Swezey); Upi, 4 9, 1 3 (2 9, 4 ¢ CAS) under bark of Pariti tiliaceum, 5.v.1936 (Swezey);
Apra, 9 2, 2 3 under dead bark, 15.vi.1936; Machaneo, 2 2 (3 9, 2 § CAS) on Elaeocarpus joga bark,
30.vi.1936 (Swezey); Piti, 1 2, 3 ¢ (492, 3 g CAS) under Heritiera littoralis bark, 7.x.1936 (Swezey); Yigo,
12(1 3 CAS) under dead bark, 1 2 (1 29 CAS) under Elaeocarpus joga bark, 18.x.1936 (Swezey); 192,24 ?
Guam [unlabelled from Floyd Andre Collection] (BMNH). [? Philippines]: Occ. Negros, 1 2 on Papaya,
1.9.1929 (W. D. Pierce) (CAS). New Guinea: Maffin Bay, | 9, vii.1944; 7 9, 1 3, viii.1944; 1 2, 1g, ix.1944;
1 9, 2 g, 3.x.1944; 12 9, 7 3 on bark, ix.1944 (E. S. Ross) (CAS). Solomon Is: 5 9, 4 3, 1965 (J. A. Grant);
Guadalcanal, Umasani River, 2 2, 1 ¢ on Areca macrocalyx, 1-9.vii.1965 (P. N. Lawrence); Kakum,
1 2, 3 g in arable soil, 1962 (P. Greenslade); Oreke, 1 3, 14.xii.1934 (R. A. Lever); North Guadalcanal,
2 2, 2 3 on dried calyx of shrivelled coconut, x.1931 (R. A. Lever); Rendova, 1 2 on Cocos trunk, 28.iii.1935
(R. A. Lever) (BMNH).
Machatothrips biuncinatus Bagnall
(Figs 53, 56, 72)
Machatothrips biuncinata Bagnall, 19085: 189. Holotype 2 (fore leg only), NEw GUINEA (BMNH)
[examined].
Adiaphorothrips simplex Bagnall, 19096 : 537. Lectotype 3, BORNEO (BMNH) [examined]. [Synonymized
by Mound, 1968: 134.]
Machatothrips montanus Priesner, 1932 : 344. Holotype 2, SARAWAK (SMF) [examined]. Syn. n.
Bagnall described Adiaphorothrips simplex from two males collected together, and Mound
(1968) designated one of these as lectotype. The holotype of montanus has the anteromarginal
setae missing, but the small setal bases indicate that they were probably short as in typical
biuncinatus. The original description states that the fore femora have only three teeth, but three
extra small humps are visible in the position of the typical fused comb of biuncinatus. This is a
variable species with very short anteromarginal setae, long postocular setae pair II, long antero-
angulars, long antennal segment III and a rounded pelta. It is similar to artocarpi, but that species
has longer anteromarginals, shorter anteroangulars and postocular setae pair II, and a pointed
pelta (Fig. 71).
SPECIMENS STUDIED.
Malaya: Lenggong, | 2 (A. M. Lea) (CAS); Damansara Forest Reserve, 8 miles [13 km] north of Kuala
Lumpur, 1 3 on dead branches and leaves, 3.x.1973; Gombak Field Station, 16 miles [26 km] east of
Kuala Lumpur, | 9, 2 ¢ on dead wood, 25.ix.1973; Gombak, on dead branches, 1 9, 30.ix.1973, 1 2, 4 3,
5.x.1973, 1 3, 8.x.1973; Ringlet, 2 2 on a dead branch, 12.x.1973 (L. A. Mound) (BMNH). Singapore:
Bukit Timah Forest, 1 9, 3 ¢ on fallen Dialium wallichii tree, 16.xi.1973 (D. H. Murphy) (BMNH).
Mentawai Is: Siberut, 1 3, 13.ix.1927 (BMNH). Java: Jasinga Forest Reserve, 1 9, 4 ¢ on a dead leafy
branch, 30.x.1973 (L. A. Mound) (BMNH); Marywattie near ? Lindanglaja Menzel, 2 2 under bark,
vii.1924 (SMF). Borneo: Sambas, lectotype 3 and paralectotype 3 of simplex, v.1890 (Th. F. Lucassen);
Sarawak, Lawas, | 2 (C. R. Wallace); West Sarawak, Quop, 1 3, iii.1914 (G. E. Bryant); Sarawak,
Matang, | 3 on dead bark, 8.xii.1913; 19, 1 d on dead tree, 17.xii.1913 (G. E. Bryant) (BMNH); Sarawak,
Mt Dulit, 3000 ft [1000 m], holotype 2 of montanus (Mjoberg) (SMF). Sulawesi (Celebes): 1 2 (de la
Saviniére) (BMNH); 1 2 without data from the Bagnall Collection (BMNH). New Guinea: holotype 9
of biuncinatus (Netherlands’ New Guinea Expedition, 1904-5); Bulolo Gorge, north of Wau, 2 $ on bark
of fallen log, 4.vi.1968 (R. Rice) (BMNH); Finschhafen, 5 9, 1 ¢ on bark, 16.iv.1944; Maffin Bay, 1 2
on bark, ix.1944 (E. S. Ross) (CAS).
Machatothrips braueri Karny
(Figs 57, 64, 73)
Machatothrips braueri Karny, 1912 : 23. Holotype 9, CAMEROUN (MNHU) [examined].
Machatothrips multidens Bagnall, 1934a : 487. Lectotype 2, GHANA (BMNH) [examined]. [Synonymized
by Mound, 1968: 134.]
Machatothrips paucidens Bagnall, 1934a : 489. Lectotype 9, GHANA (BMNH) [examined]. [Synonymized
by Mound, 1968 : 134.]
Machatothrips paucidens var. bicolorisetosus Bagnall, 1934a: 489. Lectotype 2, SIERRA LEONE (BMNH)
[examined]. [Synonymized by Mound, 1968: 134.]
190
SF Se
cS
igi tate Se
IIo SION
CER
i?
Ty ~
SS
ae we’
we
POS ES EG
Soe eS
79
Vere ee ee
. es rd
PX Meg peg: Shee em sagt og l\
oe iY Pd |
Figs 71-79 Machatothrips and Macrothrips species. 71-78 peltas, (71) Machatothrips artocarpi;
(72) M. biuncinatus; (73) M. braueri; (74) M. antennatus; (75) M. lentus; (76) M. heveae; (77) M.
haplodon; (78) Macrothrips species. 79, Machatothrips lentus, tergite III.
19]
Machatothrips braueri is very similar to heveae from Malaya. Both species have well-developed
postocular setae pair II, these setae being more than one-third as long as pair I, and the measure-
ments of antennal segment III as well as the anteromarginal setae lie within the same ranges.
The row of teeth on the fore femora of the females is also variable. Usually braueri has a row of
5 to 9 separate teeth, the most basal being the largest, whereas heveae has 4 to 7 separate teeth,
the most basal being smaller than the second (Fig. 63). There are, however, some braueri with
a small basal tubercle or tooth, but usually the two species may be separated on the length of the
posteroangular setae. These setae in braueri are long, more than half as long as the pronotum,
whereas those of heveae are short, seldom more than half and usually only one-half to one-third
as long as the pronotum. There are some specimens of braueri in the BMNH labelled paucidens
or bicolorisetosus which are smaller and have shorter, darker setae. The third antennal segment
of some of these is also very short, being only 3-0 to 3:5 times as long as broad.
Machatothrips braueri may be distinguished from haplodon, the only other species from
Africa, by its much longer postocular setae pair II, those of haplodon being less than 0-2 times as
long as pair I, and those of braueri being more than 0-3 times as long as pair I.
SPECIMENS STUDIED.
Sierra Leone: 1 2 on coconut (E. Hargreaves); Njala, lectotype 9, paralectotype ° of bicolorisetosus, also
1 2 with identical data, in tunnels of coffee twig borer, 7.i.1927 (E. Hargreaves); 1 9, 1 3 in twigs of
Bauhinia tomentosa, 1933 (E. Hargreaves); 2 2 in cassava stem (E. Hargreaves); 1 9, 1 3 on coffee,
16.x.1963 (CIE 19548); Petema, 1 ¢ under bark of Kola, 19.ii.1925 (E. Hargreaves) (BMNH). Ghana:
on cocoa, 2 3, vi.1972, 22 2, 7 3, 1972 (J. D. Majer); Kade, on cocoa, 1 9, 27.1.1971, 25 9, 13 3, i-1i.1972
(J. D. Majer); Aburi, in tunnels of cocoa stem moth borer, lectotype ° of paucidens, lectotype 2 and
paralectotype 3 of multidens, 29.x.1915; 5 2 paralectotypes, 15.xi.1915; 3 9, 1 gd in tunnels of Cacao shot
hole borer, 20.xi.1915 (W. H. Patterson); on Tapinanthus bangwensis on cocoa 1 9, 1 3, 1969, 8 2, 1 3,
1970 (P. M. Room) (BMNB). Nigeria: Ibadan, 2 2 on dead wood, 16.ix.1964; Ibadan, Moor Plantation,
3 2,3 gon yam leaves, 7.vii.1964 (B. A. Okwakpam) (BMNH). Zaire: 1 ¢ on Cacao (R. Mayne) (BMNH);
Kasai, Ngombe, 1 9, 8.xi.1921 (H. Schouteden) (SMF). Cameroun: Bascho, holotype 2 of braueri, iv.1909
(Oberlt. Bartsch. S.V.) (MMNHU).
Machatothrips braueri f. karnyi Priesner
Machatothrips braueri f. karnyi Priesner, 1932: 340. Holotype 9, CONGO (BRAZZAVILLE) (unknown)
[not examined].
This form was described from a unique female with three large and three small spines on the
fore femur. Unfortunately the type has not been traced and the original description is insufficient
to assign it to any one species.
Machatothrips celosia Moulton
Machatothrips celosia Moulton, 1928a : 325. Holotype 2, TAIWAN (Formosa) (CAS) [examined].
This species is known only from the holotype which is very similar to artocarpi. It may be dis-
tinguished by the anteromarginal setae being longer than the anteroangulars and by the longer
third antennal segment. The second specimen mentioned in the original description is indis-
tinguishable from typical artocarpi.
SPECIMEN STUDIED.
Taiwan: Kagi, holotype 2 on Celosia argenta, i.1928 (K. Toyota) (CAS).
Machatothrips corticosus Ananthakrishnan
Machatothrips corticosus Ananthakrishnan, 19725 : 443. Holotype 2, INDIA (TNA) [not examined].
Type-material of this species was not available for study but from the original description it
would appear to be most similar to indicus Ananthakrishnan. Both species were collected from
Kerala, India.
192
Machatothrips decorus sp. n.
(Figs 65, 68)
2 macroptera. Colour uniformly brown, basal three-fifths of tube dark brown to black; wings pale with
dark median longitudinal line in basal half; abdominal setae pale, yellowish, wing-retaining setae slightly
darker.
Head 1-5 times as long as broad, postocular setae pair I 2:8 times as long as postocular setae pair IT;
antennal segment III 3-6 times as long as broad. Pronotum twice as broad as long, anterior angles with a
row of three small spine-like setae; anteroangular setae twice as long as anteromarginals; median length
of pronotum 1:2 times as long as posteroangular setae. Fore femora enlarged with a row of 12 to 14
small teeth in apical half of inner margin; inner margin of fore tibiae weakly convex, fore tarsal claw small.
Tube evenly narrowing to apex, 3-3 times as long as basal breadth.
Measurements (holotype 2 in um). Body, length 5420. Head, length 500; maximum breadth across
cheeks 355. Postocular setae pair I 220; pair II 76. Antennal segments, IIT length 220, breadth 62; length,
IV 210; V 176; VI 134; VII 80; VIII 76. Pronotum, median length 270; maximum breadth 548; antero-
marginal setae 58; anteroangular setae 119; mid-lateral setae 128; posteroangular setae 214; epimeral
setae 286. Fore wing, length 2208; maximum breadth 240; number of duplicated cilia 63; length basal
setae, 1, 142; 2, 190; 3, 334. Dorsal setae tergite IX 600. Tube, length 670; maximum breadth 200.
3d macroptera. Colour and structure similar to 2. Posteroangular setae shorter than in 9, pronotum 2-4
times as long as these setae. Fore femora aberrant, with two well-developed and two small teeth on inner
margin; fore tarsal tooth sharply pointed, as long as its basal breadth (Fig. 68).
Measurements (paratype ¢ in um). Body, length 4675. Head, length 470; maximum breadth across
cheeks 316. Postocular setae pair I 190; pair II 58. Antennal segments, III length 180, breadth 58; length,
IV 176; V 162; VI 120; VII 70; VIII 70. Pronotum, median length 270; maximum breadth 524; antero-
marginal setae 48; anteroangulars 95; mid-laterals 100; posteroangulars 114; epimerals 200. Fore wing,
length 2016; maximum breadth 190; duplicated cilia 54; basal setae, 1, 90; 2, 124; 3, 152. Dorsal setae
tergite IX 476. Tube, length 546; maximum breadth 152.
SPECIMENS STUDIED.
Holotype °, Malaya: Gombak, 16 miles [26 km] west of Kuala Lumpur, on dead branches, 8.x.1973
(L. A. Mound) (BMNH).
Paratype. 1 3, same data as holotype.
This species is most similar to antennatus in the shape of the third antennal segment, the small
spine-like setae on the fore angles of the pronotum and the long row of small tubercles on the
female fore femora. It may be distinguished mainly by the much longer postocular setae pair II.
It is also not easily distinguishable from the African species braueri, apart from the arrangement
of the fore femoral teeth of the female and slightly shorter anteromarginal setae. The male is
unusual in having teeth on the fore femora.
The two specimens on which this species is based were collected with a male of biuncinatus.
Machatothrips diabolus (Priesner)
Adiaphorothrips diabolus Priesner, 1928a : 56. Holotype 2, EAst AFRICA (Hamburg Museum, destroyed).
This species was described from a unique female which was destroyed in the Hamburg Museum.
It was not illustrated and the original description is insufficient even to assign it to the genus
Machatothrips. Priesner (1928a) states that it most closely resembles e/ephas Karny which he
subsequently transferred to the genus Scotothrips. The name diabolus is best regarded as a nomen
dubium at present.
Machatothrips haplodon Karny
(Figs 60, 77)
Machatothrips braueri var. haplodon Karny, 1925: 141. Holotype 2, UGANDA (BMNH) [examined].
Machatothrips braueri var. buffai Karny, 1925 : 142. Holotype 3, UGANDA (BMNH) [examined]. [Synony-
mized by Mound, 1968 : 135.]
Machatotothrips simplicidens Bagnall, 1934a: 490. Holotype 2, CoNGO (BRAZZAVILLE) (MNHN) [not
examined]. [Synonymized by Mound, 1968 : 135.]
193
Machatothrips braueri f. spatiata Priesner, 1932: 341. LECTOTYPE 92, Zarre (SMF), here designated
[examined]. Syn. n.
Machatothrips haplodon Karny; Mound, 1968 : 135.
Machatothrips braueri f. spatiata was described for an unspecified number of female syntypes
from various localities in the Belgian Congo. Three females labelled ‘paratype’ have been studied.
One, from Kasai, Ngombe, 8.xi.1921 (Dr H. Schouteden), has postocular setae pair II 0-3 times
as long as pair I, antennal segment III 4-0 times as long as broad, and a short blunt fore tarsal
claw. The fore femoral teeth are variable in form and the presence of a small tooth at the base is
not sufficient to separate this specimen from typical braueri. The two other ‘paratypes’, one from
Haut Uelé, Moto, 1920 (L. Burgeon), and the other from Barumbu here designated as lectotype,
have postocular setae pair II 0-1 times as long as pair I, antennal segment III 3-2 to 3-5 times as
long as broad and a long and sharply pointed fore tarsal claw. The presence of the small spine
at the base of the femora of these two specimens is not sufficient to separate them from typical
haplodon. Machatothrips braueri var. buffai was described from specimens without fore femoral
teeth. The holotype is a male which, in common with most males of this genus, has no fore
femoral teeth. It has short postocular setae pair II and is indistinguishable from typical haplodon.
SPECIMENS STUDIED.
Ghana: 1 2 on Tapinanthus banguensis on cocoa, 1970 (P. M. Room) (BMNH). Zaire: lectotype 2 of
spatiata, Barumbu, viii.1925 (J. Ghesquiére), paralectotype 2 of spatiata, Haut Uelé, Moto, 1920 (L.
Burgeon) (SMF). Uganda: Kampala, holotype 2 of haplodon, holotype 3 of buffai and 2 2 in Scolytid
galleries in bark of Ficus ovata, 8.viii.1921 (H. Hargreaves) (BMNH).
Machatothrips heveae Karny
(Figs 58, 63, 76)
Machatothrips heveae Karny, 1921b: 61. Holotype 2, JavA (SMF) [examined].
This species is very similar to the African species braueri but may usually be distinguished by its
shorter posteroangular setae.
SPECIMENS STUDIED.
Malaya: Kuala Lumpur, 2 2 (A. M. Lea) (CAS), 2 2, 3 3 on dead branches, 29.xii.1969 (R. G. & F.
Andre); Gombak, 2 3 on dead branches 29.ix.1973, 1 2 on Cassia leaves, 2.x.1973 (L. A. Mound) (BMNH).
Java: Halte Dawoean, holotype 2 on Hevea xi.1906 (De Vos V.N.C.) (SMF).
Machatothrips indicus Ananthakrishnan & Jagadish
(Fig. 59)
Machatothrips indicus Ananthakrishnan & Jagadish, 1970: 279. Holotype 2, INDIA (TNA) [examined].
This species, described from two females and two males, may be distinguished easily from the
others in the genus by the inner margin of the fore tibiae being set with small spinules in both sexes
and the fore tibiae of the females having a distinct hump.
SPECIMENS STUDIED.
India: Aryankavu, holotype 2 and paratype ¢ on dry twigs, 18.vii.1969 (TNA).
Machatothrips lentus sp. n.
(Figs 61, 69, 70, 75, 79)
2 macroptera. Colour dark brown to black, basal stem of antennal segment III, tarsi and extreme apex
of tube slightly paler. Wings dusky with a dark median longitudinal line in basal half. Wing-retaining
setae dark brown, the two pairs of tergal posteroangular setae pale, colourless to yellow.
Head about 1-5 times as long as broad, postocular setae pair I 3-0 to 6:5 times as long as pair II.
Antennal segment III 3-2 to 3-8 times as long as broad. Pronotum twice as broad as long, anteroangular
194
setae 1-5 to twice as long as anteromarginals, median length of pronotum 1-7 to 2:5 times as long as the
short posteroangulars. Fore femora enlarged with 2 to 4 separate teeth plus a ridge of 3 to 6 in the distal
half of the inner margin; inner margin of fore tibiae with a distinct hump opposing the basal, largest,
femoral tooth; fore tarsal claw small. Tube evenly narrowing to apex, about 3-5 times as long as broad.
Measurements (holotype 2, also largest and smallest 2 paratypes, in um). Body, length 5328 (5874,
5106). Head, length 520 (546, 500); maximum breadth across cheeks 326 (344, 316). Postocular setae,
pair I 200 (209, 186), pair II 66 (62, 52). Antennal segments, III length 200 (214, 190), breadth 62 (62, 58);
length, IV 186 (204, 180); V 156 (170, 158); VI 120 (128, 120); VII 76 (76, 76); VIII 76 (80, 76). Prono-
tum, median length 266 (314, 268); maximum breadth 530 (540, 510); anteromarginal setae 42 (62, 42);
anteroangulars 66 (80, 62); mid-laterals 90 (95, 86); posteroangulars 120 (132, 110); epimerals 228 (248,
214). Fore wing, length 2160 (2400, 2062); maximum breadth 220 (240, 200); duplicated cilia 57 (67, 60);
basal setae, 1, 90 (120, 90); 2, 148 (200, 148); 3, 248 (286, 238). Dorsal setae tergite IX 575 (592, 564).
Tube, length 632 (710, 632); maximum breadth 180 (190, 180).
3 macroptera. Colour and structure similar to 9. Inner margin of fore femora normally without teeth,
inner margin of fore tibiae without hump; fore tarsi with well-developed, rounded claw. Some males
have a tendency to produce fore femoral spines; these specimens have shorter, more sharply pointed,
fore tarsal claws.
Measurements (largest and smallest 3 paratypes in um). Body, length 4942, 4800. Head, length 490,
480; maximum breadth across cheeks 306, 306. Postocular setae, pair I 190, 186; pair II 28, 42. Antennal
segments, III length 200, 194, breadth 58, 58; length, IV 186, 180; V 162, 156; VI 122, 120; VII 70, 70;
VIII 70, 70. Pronotum, median length 366, 304; maximum breadth 592, 528; anteromarginal setae 52,
40; anteroangulars 80, 80; mid-laterals 114, 104; posteroangulars 134, 114; epimerals 238, 224. Fore
wing, length 2160, 2015; maximum breadth 214, 200; duplicated cilia 74, 63; basal setae, 1, 90, 86; 2,
142, 138; 3, 228, 214. Dorsal setae tergite IX 466, 472. Tube, length 584, 536; maximum breadth 162, 162,
SPECIMENS STUDIED.
Holotype 2, Malaya: Genting Highlands, 4500 ft [1500 km], 30 miles [48 km] east of Kuala Lumpur,
dead wood and leaves, 28.ix.1973 (L. A. Mound) (BMNH).
Paratypes. Malaya: Damansara Forest Reserve, 8 miles [13 km] north of Kuala Lumpur, | 2, 3 3 on
dead branches and leaves, 3.x.1973; Tanah Rata, 3 2, 1 g on dead branch, 12.x.1973; Ringlet, 1 2, 1 g on
dead branch, 12.x.1973; Gombak, on dead branch, 1 9, 5.x.1973, 1 3, 13.x.1973 (L. A. Mound); Cameron
Highlands, 1 2 under bark of Citrus, 8.xi.1974 (J. A. Whellan) (BMNH); Kuala Lumpur 1 3; Fraser’s
Hill, Gap, 2 2 (A. M. Lea) (CAS). Singapore: 1 2 (A. M. Lea & C. T. McN.) (CAS).
This species is most similar in colour and form to artocarpi but it may usually be distinguished
by the shorter third antennal segment, posteroangular setae and postocular setae pair II. The
females, however, are distinctive in having a hump on the inner margin of the fore tibiae. The
teeth on the fore femora are arranged similarly to biuncinatus with the smaller apical teeth some-
times fused at the bases to form a ridge. This species appears twice in the key due to the variation
in length of the postocular setae pair II, pair I being 3-0 to 6-5 times as long as these setae.
Machatothrips quadrudentatus Moulton
(Fig. 55)
Machatothrips quadrudentatus Moulton, 1947 : 179. Holotype 2, NEw GuINEA (CAS) [examined].
This species is known only from the holotype which, although described as a male, is, in fact, a
female. It is distinct in having a short pale, third antennal segment 2:5 times as long as broad, very
short postocular setae pair II, which are only 0-1 times as long as pair I, and long anteromarginal
setae equal in length to the anteroangulars. The fore femora bear four simple separate teeth in
the distal half.
SPECIMEN STUDIED.
New Guinea: Finschhafen, holotype 9, v.1944 (E. Ross) (CAS).
Machatothrips silvaticus Ananthakrishnan
Machatothrips silvaticus Ananthakrishnan, 1972a : 436. Holotype 2, INDIA (TNA) [paratypes examined].
This species is known from 21 male and 35 female paratypes, in addition to the holotype, from
195
India. It has a short, third antennal segment, slightly paler than the fourth and may be dis-
tinguished from indicus mainly by the lack of tubercles on the fore tibiae.
SPECIMENS STUDIED.
India: Madhya Pradesh, Kanha forest, 2 g, 1 2 paratypes on dry twigs, 21.i.1970 (TNA).
MALESTIA THRIPS gen. n.
Type-species: Malesiathrips malayensis sp. n.
Small, dark or bicoloured species of Idolothripinae. Head longer than wide, dorsal surface sculptured,
bearing several pairs of broadly blunt, pale setae which arise from tubercles, maxillary stylets V-shaped.
Antennae 8-segmented, III-IV less than twice as long as wide; 2 sense cones on III, 4 sense cones on IV.
Pronotum with epimeral sutures complete; 5 pairs of major setae present, anteroangular and midlateral
setae relatively close together on a slight ridge, epimeral setae elongate on a small tubercle; praepectal
plate broad, parallel to probasisternum. Fore tarsus with or without a tooth; all femora with one or more
stout pale setae. Mesonotal lateral setae small; metanotum reticulate, median setae stout. Fore wings
slightly angled before middle, parallel-sided distally, without duplicated cilia; sub-basal setae B, stout.
Pelta broad; abdominal tergites with anterior pair of wing-retaining setae small or absent; posterior
margin of tergites with 4 pairs of setae; tergite IX setae B, and B, stout, B, finely acute; tube less than 4
times as long as tergite IX, terminal setae short.
This new genus is erected for three species from the Malesian phytogeographic region. One of
these species, solomoni Mound, was described in the genus Atractothrips but the type-species of
that genus, from Florida, has the following characteristics: antennal segment III more than 4
times as long as wide, sense cones less than 0:25 times as long as segment; praepectus absent;
pronotal midlateral setae reduced; sub-basal wing setae not developed; tergites with 2 pairs of
sigmoid wing-retaining setae; posterior margin of tergites with 3 pairs of setae; tergites sculptured
in front of antecostal ridge; tube more than 5 times as long as tergite IX.
Malesiathrips appears to be similar to the monobasic Hawaiian genus Dermothrips which has
very reduced setae and the terminal antennal segments condensed. However, the tribal relation-
ships of these genera, together with Emprosthiothrips from Australia, are at present uncertain.
Key to species
1 Antennal segment II bearing 1 large seta, with expanded apex, which extends beyond mid-point
of III (Fig. 82); head prolongation in front of eyes shorter than diameter of one ocellus;
head with 1 pair of postocular setae (Fig. 80); sub-basal setae shorter than half basal width
of fore wing; median metanotal setae short, about as long as one sculptured reticle [Guam]
guamensis (p. 196)
— Antennal segment II with 2 short broad setae at apex (Fig. 83); head prolongation as long as
diameter of one ocellus; head with 2 pairs of postocular setae (Fig. 81); sub-basal setae longer
than half basal width of fore wing; median metanotal setae longer than the diameter of 2
reticles . : A ; ; ; : ; ; : : : R : : 2
2 Head with 1 pair of setae on vertex behind and equal in length to postocular setae; median
area of vertex tuberculate; pronotal posteromedian discal setae short and acute; fore wing
with 2 long sub-basal setae; 2 with small fore tarsal tooth; head dark laterally, antennal
segments III-IV light brown, legs bicoloured, tube as dark as abdomen [Solomon Is]
solomoni (p. 199)
— Head with 2 or 3 pairs of short broad setae behind postoculars (Fig. 81); median area of
vertex with large, transverse reticles; pronotal posteromedian discal setae as long as antero-
marginals; fore wing with 1 long sub-basal seta; 2 without a fore tarsal tooth; head with 2
longitudinal pale stripes, antennae blackish brown except pedicel of III, legs uniformly dark,
tube much paler than abdomen [Malaya, Java]. ‘ ‘ : , malayensis (p. 198)
Malesiathrips guamensis sp. n.
(Figs 80, 82)
Macropterous 9. Colour brown; abdomen and tube, pronotum and sides of pterothorax dark brown;
head, median area of pterothorax and pelta light brown, legs dark, tarsi yellow; antennal segments,
196
Figs 80-85 Malesiathri, i
ips species. (80) M. guamensis; (81) M. mala is; j
: : yensis; (82) M. :
(83) M. maiayensis; (84) M. malayensis, pelta; (85) M. malayensis, ae oes
197
I-II, VI-VIL dark, III-IV mainly yellow, V yellow to brown; fore wing weakly shaded medially; major
setae pale, but large dorsal seta on antennal segment II dark.
Head with irregular tubercles medially, reticulate between eyes and at posterior margin; eyes rather
small; postocular and postocellar setae little larger than cheek setae. Antennal segment II with median
dorsal seta long and spatulate; III-IV with dorsal setae expanded at apex; pedicel of VI broad. Pronotum
sculptured; major setae almost tubular. Metanotum strongly reticulate, median setae small. Pelta reti-
culate, broad, close to anterior margin of tergite II; tergites similar to malayensis (Fig. 85); tergite IX
with setae B, and B, broadly rounded at apex, B, finely acute; tube margins weakly convex, with longi-
tudinal ridges basally.
Measurements (holotype 2 in um). Body, length (contracted) 2000. Head, length 325; median width
220; postocellar and postocular setae 25. Pronotum, length 180; median width 300; major setae, antero-
marginal 15, anteroangular 30, mid-lateral 17, epimeral 70, posteroangular ? 30. Metanotal median
setae 15. Fore wing, length 1000; distal width 80; sub-basal setae 12, 15, 30. Tergite IX setae, B,, 75;
B,, 75; Bs, 90. Tube, length 300; terminal setae 150. Antennal segments, length, 45, 70, 75, 72, 60, 45,
40, 30.
SPECIMEN STUDIED.
Holotype 2, Guam: Upi, under bark of Pariti tiliaceum, 5.v.1936 (Swezey) (BMNH).
The eyes of guamensis appear to be more reduced than in the other two species of the genus.
The pedicillate antennal segments are similar to solomoni, but there is a large seta on segment
II. The tergites appear to be similar to malayensis, but the abdomen of the unique holotype is
contracted and difficult to study.
Malesiathrips malayensis sp. n.
(Figs 81, 83-85)
Macropterous 9. Colour yellow to blackish brown; head pale between the eyes and with two submedian
pale stripes, rest of head dark; antennal segments II-VIII dark except base of III; thorax and legs brown,
tarsi paler; abdominal segments II-VUI blackish brown, slightly paler submedially, LX pale, tube reddish
brown; fore wings shaded, particularly at base, with a longitudinal dark stripe.
Head projecting in front of large eyes; dorsal surface reticulate medially, tuberculate laterally (Fig. 81);
postocellar and lateral postocular setae sometimes longer than median postocular setae. Antennal
segments [V—VI with broad pedicels; dorsal setae on I-V flattened or apically blunt. Pronotum with
posteromedian discal setae similar in structure to the 5 pairs of major setae. Tergites with median pores
close together, III-VIII with anterior pair of wing-retaining setae weakly sigmoidal, II—-VII with posterior
pair of wing-retaining setae sigmoid and flattened (Fig. 85); sternal marginal setae very small; tube
parallel-sided, ridged near base, constricted at apex.
Measurements (holotype 2 in um). Body, length (extended) 2400. Head, length 345; median width
185; postocellar setae 65; postocular setae 30, 65. Pronotum, length 135; median width 255; major setae,
anteromarginal 35, anteroangular 54, mid-lateral 30, epimeral 105, posteroangular 54. Metanotal median
setae 54. Fore wing, length 1000; distal width 75; sub-basal setae 2, 10, 65. Tergite IX setae, B,, 110;
B;, 95; Bz, 80. Tube, length 330; terminal setae 90. Antennal segments, length 50, 60, 75, 68, 68, 50,
40, 30.
Apterous 3. Similar in colour and structure to 2; head without ocelli; fore tarsus with a small, broad-
based tooth; tergites with 2 pairs of large posteromarginal setae and 2 pairs of minute setae; tergal postero-
angular setae small but thorn-like.
Measurements (paratype ¢ from Java, in um). Body, length 1800. Head, length 270; median width 165;
postocellar setae 50; postocular setae 30, 48. Pronotum, length 125; median width 225; epimeral setae 60.
Tube, length 210. Antennal segments, length, 50, 50, 65, 50, 50, 40, 35, 25.
SPECIMENS STUDIED.
Holotype 9, Malaya: 16 ml [26 km] east of Kuala Lumpur, Gombak Field Station, on dead wood,
26.ix.1973 (L. A. Mound) (BMNB).
Paratypes. Malaya: Kuala Lumpur, 3 2 on dead leaves and branches, 23—27.xii.1969 (R. G. & F.
Andre) (BMNH). Java: Bogor Botanic Gardens; 30 2, 8 ¢ from dead palm fronds, 24.x.1973 (L. A.
Mound); 4 2 from dead leaves, 25.x.1973 (L. A. Mound) (BMNHB).
198
The head of malayensis is similar in shape to that of solomoni, but the antennal segments are
more condensed, and the fore tarsal tooth is smaller. Moreover, malayensis has 2 pairs of wing-
retaining setae whereas the anterior pair is not developed in solomoni. The paratypes from Java
and Malaya have shorter setae on the head than the holotype.
Malesiathrips solomoni (Mound) comb. n.
Atractothrips solomoni Mound, 1970: 116-118. Holotype 9, SoLoMoN Is (BMNH) [examined].
This species is discussed under guamensis and malayensis. The male has the tergal posteroangular
setae short but stout and thorn-like.
SPECIMENS STUDIED.
Solomon Is: Choiseul I., Vasu River, holotype 2 with 2 9, 1 ¢ paratypes from forest litter, 16.xi.1965
(Greenslade) (BMNH).
MECYNOTHRIPS Bagnall
Mecynothrips Bagnall, 1908a : 356. Type-species: Mecynothrips wallacei Bagnall, by original designation
and monotypy.
Phoxothrips Karny, 1913 : 132. Type-species: Phoxothrips pugilator Karny, by monotypy. Syn. n.
Kleothrips Schmutz, 1913 : 1057. Type-species: K/eothrips gigans Schmutz, by monotypy, here regarded
as a synonym of Mecynothrips simplex Bagnall, 1912. Syn. n.
Dracothrips Bagnall, 1914: 290. Type-species: Dracothrips ceylonicus Bagnall, by monotypy, here re-
garded as a synonym of Mecynothrips simplex Bagnall, 1912. Syn. n.
Acrothrips Karny, 1921b: 43. Type-species: Acrothrips sorex Karny, by monotypy, emended from serex
by Karny, 1924: 37, here regarded as a synonym of K/eothrips acanthus Hood, 1918. Syn. n.
Kleothrips (Synkleothrips) Priesner, 1935c : 330. Type-species: K/eothrips (Synkleothrips) innocens Priesner,
by monotypy, here regarded as a synonym of Kleothrips lacerta Priesner, 1935c. Syn. n.
Kleothrips (Akleothrips) Priesner, 1935c: 332. Type-species: Kleothrips (Akleothrips) karimonensis
Priesner, by original designation. Syn. n.
Mecynothrips Bagnall; Mound, 19715 : 279-283.
Phoxothrips Karny; Haga & Okajima, 1974 : 375-384.
Large, macropterous species of Idolothripini. Head elongate, 2 to 3 times as long as greatest breadth,
prolongation in front of eyes also elongate, 2 to 3 times as long as broad; eyes sometimes slightly longer
on the dorsal surface; head with 2 pairs of elongate ocellar setae, pair I near first ocellus, pair II posterior
to ocellar triangle; 1 pair of postocular setae and 1 pair of median setae on vertex; cheeks with at least 3
(usually more than 6) pairs of pale, stout spine-like setae arising from tubercles. Antennae 8-segmented,
segment III 6-0 to 11-5 times as long as broad with 2 sense cones, IV with 4 sense cones. Pronotum about
one-third as long as total head length, 1-5 times as broad as long to slightly longer than broad, usually
with 5 pairs of major setae; anteromarginals, anteroangulars and mid-laterals sometimes reduced to
minor setae, and the second pair of epimerals sometimes well developed ; fore angles of some males bearing
large recurved horns; epimeral sutures sometimes not developed in larger males. Fore femora of males
and females enlarged, rectangular or bulbous in males, usually with an apical tooth and with or without
a median or basal tubercle. Fore tibiae of males sometimes with a seta-bearing tubercle at apex; fore
tarsal claw well developed in most males, absent in females and some gynaecoid males. Wings broad,
slightly broader in apical half, pale or dusky with numerous duplicated cilia. Pelta broad with lateral
lobes. Tergite II with at least 2 pairs of wing-retaining setae, tergites III-V with 3 pairs of sigmoid wing-
retaining setae and numerous sigmoid accessory setae. Sternite IX of males with a pair of stout setae.
Median pair of setae on tergite IX 0-40 to 1-25 times as long as tube. Tube with straight sides evenly
narrowing to apex, 4 to 5 times as long as broad, 1:10 to 3-75 times as long as head prolongation, with
a few small setae laterally.
The length of the head prolongation is measured laterally from the base of the antenna to the
anterior margin of the eye, and the breadth is measured across the base anterior to the eyes.
The ‘head length from behind eyes’ is also measured laterally between the posterior margin of
the eyes and the posterior margin of the vertex.
199
The monotypic genus Dracothrips was erected by Bagnall for cey/onicus, but he later synony-
mized this with Kleothrips gigans. The monotypic genus Acrothrips was erected by Karny for
sorex, but Bagnall later synonymized this with Kleothrips acanthus. Priesner erected two sub-
genera of Kleothrips; Synkleothrips for his species innocens which has unarmed fore femora and
Akleothrips for his species karimonensis which has an apical tooth and a basal tubercle on the fore
femora. Haga & Okajima (1974) indicated that the two genera Phoxothrips and Kleothrips could
not be separated and that Phoxothrips was the oldest name. Moreover, they illustrated clearly the
range in form of the male fore femur of the type-species pugilator which varies from being slender,
unarmed and female-like to being enlarged, with a median hump and apical tubercle. Despite
this variation within one species, they retained the subgeneric names Synkleothrips and Akleo-
thrips with the resulting anomaly that large and small males of pugilator would be placed in
separate subgenera. Similar variation in the fore femora occurs in hardyi Priesner (Mound, 1974)
and it appears that only one generic name is useful for this group of species.
The genus Mecynothrips was erected by Bagnall for wallacei which was based on one large
male. The distinguishing characteristics of this genus appear to be the long head prolongation,
the horns on the fore angles of the pronotum and the relatively short tube. The fore femora of
the males have an apical tooth and a median tubercle. The pronotal horns, even in small males,
are well developed and all the male specimens examined have at least a small median tubercle
on the fore femora. However, acanthus is intermediate between Phoxothrips and Mecynothrips
in the form of the fore angles of the pronotum, and the species assigned to Mecynothrips appear
to represent the more extreme oedymerous forms of Phoxothrips. The present authors do not
regard these differences as sufficient to segregate two genera in which, moreover, the females
are not easily distinguishable even at species level. Therefore Phoxothrips and Mecynothrips are
here synonymized.
Within Mecynothrips three species-groups can be distinguished on characters which are avail-
able in both sexes. In the simplex-group, which includes atratus, goliath and pugilator, the median
setae on tergite IX are shorter than the tube. In the acanthus-group, which includes hardyi,
karimonensis and lacerta, the median setae on tergite [IX are longer than the tube and the pronotal
anteromarginal setae are well developed. In the wallacei-group, which includes kraussi, priesneri
and snodgrassi, the median setae on tergite IX are longer than the tube but the pronotal antero-
marginals are small, minor setae. These species-groups appear to be correlated with the geo-
graphical distribution of the species each includes. The simplex-group is the most northern:
East Africa, Sri Lanka, Java, Sarawak, Philippines, Taiwan, Japan. The wallacei-group is the
most eastern: New Guinea, Australia, Solomon Is. The acanthus-group is the most southern:
Java, Sumba, Australia. The genus Mecynothrips appears to be the Austro-Oriental equivalent
of the otherwise pantropical genus Elaphrothrips (Mound, 1974). In both sexes it may be dis-
tinguished by the presence of 3 pairs of wing-retaining setae on tergites III-V whereas species of
Elaphrothrips have only 2 pairs.
Key to species
1 Median pair of setae on tergite IX (B,) shorter than tube, 0-4 to 0-8 times as long
— Median pair of setae on tergite IX (B,) longer than tube, 1:05 to 1-25 times as long
2 Median length of pronotum‘of 2 1-7 to 2:5, of 3 2:4 to 2:7 times as long as anteroangular setae:
legs bicoloured; fore femur of large ¢ without any development of the inner margin, apart
from a black apical spine . 3
— Median length of pronotum of 2 3-0 to 3-3, of 3 3: 9 to 7: af times as s long as anteroangular
setae; legs usually dark; fore femur of large 3 with a black apical spine and a basal bulge
or median tubercle . 4
3 Antennal segment III with a dusky basal stem, IV and Vv with a ‘darker stem and pale median
band, VI dark; fore wings with a dark median longitudinal bar at the base; 3 with a slim,
curved fore tarsal tooth and simple fore tibia (Fig. 103) [Sumba] . ‘ . goliath (p. 203)
— Antennal segments III-IV with a pale or slightly dusky basal stem; fore wings with a pale
basal half and dark margins in the apical half; 3 with long, slim, straight fore tarsal tooth;
small 3 with simple fore tibia, larger 3 with two ae nee at the base (Fig. 99)
[India to Philippines] : , ' ; : ; : . simplex (p. 207)
200
Un
, M. atratus
, M. simplex, tergite IV. 91, 92, smallest males; (91) M. kraussi; (92) M.
201
90
snodgrassi. 93, M. atratus, pelta.
Figs 86-93 Mecynothrips species. 86, M. pugilator; 87, M. acanthus; 88, M. kraussi,; 89
(zuluensis paratype);
Setae B, on tergite IX short, no longer than 0-5 times as long as tube; all legs and tarsi dark;
antennal segments III-V with a pale basal stem, segment VI completely dark; 3 with a
slim fore tarsal tooth, fore femur with a black apical spine and median tubercle (Fig. 102).
Head length from behind eyes 2-4 to 2:7 times as long as the prolongation (Fig. 89) [East
and South Africa] . ; . atratus (p. 203)
Setae B, on tergite IX longer, 0-5 to 0-7 times as slong: as tube, all legs dark, tarsi paler, antennal
segment III-VI with a pale basal stem. Large ¢ with a slim fore tarsal tooth, fore femur
with a black apical spine and basal bulge, small 3 with only a small fore tarsal tooth and
fore femur without a basal bulge. Head length from behind eyes 3-5 to 4-4 (5-5 in takahashii)
times as long as the prolongation (Fig. 86) [Japan, Taiwan] . 4 pugilator (p. 207)
Anteromarginal setae well developed, 0-75 to 1-0 of anteroangular setae (Fig. 113); tendency
to relatively longer tube and shorter head prolongation; 3 with tube more than 2-5 times
length of head prolongation; 3 without process over tentorial pits on ventral surface of head 6
Anteromarginal setae small, hardly more developed than other minor pronotal setae; tendency
to relatively shorter tube and longer head prolongation; 3 with tube less than 2:5 times
length of head prolongation; ¢ with process over tentorial pits on ventral surface of head
(Figs 88, 106, 107) . : 9
Antennal segment III dark, only slightly paler at extreme base, segments IV and Vv with 2 a
pale basal stem and dark basal ring; head prolongation about 0-25 times length of head from
behind eyes. Major 3 with a small fore tarsal tooth, also fore femur with a black apical
spine and a basal tubercle, small 3 with unarmed fore legs [Australia] . 2 hardyi (p. 203)
At least antennal segments III-V with a pale yellow basal stem; head prolongation about
0-33 times length of head from behind eyes. 3
Mid and hind tibiae not distinctly bicoloured; basal stem of antennal segment Ill paler than
IV-VI; 3 with a small, slim fore tarsal tooth and unarmed fore femur (Fig. 101) [Sumba]
lacerta (p. 206)
Mid and hind tibiae distinctly bicoloured; basal stem of antennal segment III concolourous
with IV and V. : ; 8
Basal stem of antennal segments I-VI pale yellow: 3 with a well- developed, slim fore tarsal
tooth, fore femur with an apical spine and basal tubercle (Fig. 100) [Java] karimonensis (p. 205)
Antennal segments III—-V with a dusky basal stem and a paler median band; male with a stout
fore tarsal tooth, fore femur with two apical spines but without any basal tubercle (Fig. 98)
[Australia] ; . acanthus (p. 202)
Cheeks with only 1 pair of spine- aR setae in basal two-thirds (Figs 88. 91); median sub-basal
wing seta usually longer than the most basal; 3 with cluster of 3 to 5 pairs of spine-like
setae on cheeks just posterior to eyes, fore tarsus with a long retroussé claw 3 to 4 times as
long as its basal breadth (Fig. 94); process over tentorial pits bilobed (Fig. 106) [Solomon Is]
kraussi (p. 205)
Cheeks set with numerous spine-like setae throughout entire length (Fig. 92); sub-basal wing
setae 1 and 2 subequal, or 2 shorter than 1; fore tibia of ¢ with straight fore tarsal claw,
usually only 2 to 3 times as long as basal breadth (Figs 95,96) . ‘ 10
Male fore tarsus with a stout claw about 2:5 times as long as basal breadth; fore ocellus situated
anterior to ocellar setae (Fig. 92); process over tentorial pits bilobed [Solomon Is]
snodgrassi (p. 208)
Male fore ocellus situated lateral or posterior to ocellar setae; process over tentorial pits with ;
llobe . :
Male fore tarsal claw stout, only about twice as slong as basal breadth; seta bearing tubercle at
apex of fore tibia not so distinct (Fig. 96) [New Guinea, New Britain] : priesneri (p. 207)
Male fore tarsal claw more slender, more than 3 times as long as basal breadth; seta-bearing
tubercle at apex of fore tibia distinct (Fig. 95) [New Guinea, Australia] . wallacei (p. 208)
Mecynothrips acanthus (Hood) comb. n.
(Figs 87, 98)
Kleothrips acanthus Hood, 1918: 77. Holotype 3, AUSTRALIA (USNM) [not examined].
Acrothrips serex Karny, 1920c : 43. Lectotype 3, AUSTRALIA (SMF) [examined]. [Synonymized by Bagnall,
1932 : 520.]
Acrothrips gargantua Girault, 1926: 1. Lectotype 3, AUSTRALIA (QM) [not examined]. [Synonymized
by Mound, 1974: 65.]
202
Phoxothrips giganteus Girault, 1926: 4. Holotype 3, AUSTRALIA (QM) [not examined]. [Synonymized
by Mound, 1974: 65.]
This species belongs to the group with the median setae on tergite IX longer than the tube,
well-developed anteromarginal setae and males with bulbous fore femur with a spine only at the
apex. The males may be distinguished by the presence of small horns on the fore angles of the
pronotum and a second tooth on the ventral surface of the fore femora at the apex. The females
are difficult to distinguish from karimonensis but tend to have shorter anteroangular setae and
a longer head production.
SPECIMENS STUDIED.
Australia: [Queensland] Atherton, 3 lectotype, 2 2 paralectotypes of sorex, v.1913 (Mjéberg 2) (SMF);
Kuranda 1100 ft [330 m], 2 3, 2 9, v.vi.1913, 1 3, 21.vi.—-24.vii.1913 (R. E. Turner); Kuranda—Mareeba,
Clohesy River, 1 9, 17.1.1962 (E. Britten) (BMNH).
Mecynothrips atratus (Hood) comb. n.
(Figs 89, 93, 102, 114)
Kleothrips atratus Hood, 1919: 69. Holotype 2, East AFRICA (USNM) [examined].
Kleothrips (Akleothrips) zuluensis Jacot-Guillarmod, 1939: 70. Holotype 3, SouTH ArFRIcA (JG) [ex-
amined]. Syn. n.
This species is the only representative of Mecynothrips in Africa and belongs to the group with
the median setae on tergite IX shorter than the tube. It has a distinctively shaped head and pelta
(Figs 89, 93), rather short antennal segments, incomplete epimeral sutures, pale wings and
completely dark legs. The anterior pair of wing-retaining setae on tergites III-V are sometimes
weakly developed, particularly in the female. The males have an enlarged fore femur with an
apical tooth and a well-developed median tubercle, and a slim fore tarsal claw. Kleothrips atratus
was described from one female only and zuluensis from two males. Jacot-Guillarmod distinguished
them on the shape of the head and its prolongation, also the relative lengths of antennal segments
VII and VIII and abdominal segments VIII and XI. These differences are so small that they are
here regarded as sexual variation and the two species synonymized.
SPECIMENS STUDIED.
East Africa [Tanzania]: Rufigi River, 2 holotype of atratus, 1.vi.1917 (A. W. Robbins-Pomeroy) (USNM).
South Africa (Zululand): ¢ holotype of zuluensis, Hluhluwe, Hhabisa, 24.i.1937 (Jacot-Guillarmod),
3 paratype of zuluensis, White Umfolosi, 1922 (R. H. T. P. Harris) (JG).
Mecynothrips goliath (Priesner) comb. n.
(Fig. 103)
Kleothrips goliath Priesner, 1935c : 327. Holotype 3, SUMBA (SMF) [examined].
This species is very similar to simplex in having short setae on tergite IX and unarmed, bulbous
fore femora. It may be distinguished by the darker basal stem of antennal segments III-VI, the
dark median bar at the base of the fore wings, the dark postocular setae, also the slim curved
fore tarsal claw and the unarmed fore tibiae of the males.
SPECIMENS STUDIED.
Sumba: Kananggar, 700 m, holotype 3, paratype 2, v.1925 (Dammerman 224) (SMF).
Mecynothrips hardyi (Priesner) comb. n.
(Figs 97, 113)
Kleothrips hardyi Priesner, 1928) : 657. Holotype 2, AUSTRALIA (SMF) [examined].
Kleothrips hardyi Priesner; Mound, 1974: 65-66.
This is a dark species belonging to the group which has the median setae on tergite IX longer
203
Figs 94-107 Males of Mecynothrips species. 94-97, fore tibiae and tarsi, (94) M. kraussi; (95) M.
wallacei; (96) M. priesneri; (97) M. hardyi. 98-105, fore legs, (98) M. acanthus; (99) M. simplex;
(100) M. karimonensis; (101) M. lacerta (innocens holotype); (102) M. atratus (zuluensis paratype) ;
(103) M. goliath; (104, 105) M. priesneri (104, minor paratype). 106, 107, tentorial pit covers,
(106) M. kraussi; (107) M. priesneri.
than the tube and well-developed pronotal anteromarginal setae. Large males have bulbous
fore femora with an apical tooth as well as a basal tubercle and a small fore tarsal claw. Small
males, however, have completely unarmed fore legs. This species is distinct in having a dark
antennal segment III, dark legs and dark wings in both sexes.
SPECIMENS STUDIED.
Australia: [Queensland] Brisbane Bot[anical] Gardens, holotype 2 (Hardy) (SMF); 30 ml [48 km], N.
Biloela, 4 3, 2 2 on dead Eucalyptus leaves, 18.vii.1968 (L. A. Mound) (BMNH).
Mecynothrips kanoi Takahashi comb. n.
Kleothrips kanoi Takahashi, 1937 : 343. Syntypes 2 2, TAIWAN (depositary unknown) [not examined].
This species was described from two female syntypes but has not been recognized since. From
the original description it can be established that kanoi belongs to the group which has the median
setae on tergite [X shorter than the tube. It has a shorter head prolongation than simplex, dark
legs and antennal segments [II-VI with a pale base. Takahashi states that it differs from takahashii
in the leg colour, the head prolongation and the shorter anteroangulars. It is likely, however,
that these characteristics fall within the variation of pugilator of which it is probably a synonym.
Mecynothrips karimonensis (Priesner) comb. n.
(Fig. 100)
Kleothrips (Akleothrips) karimonensis Priesner, 1935c : 332. Holotype 3, JAVA (SMF) [examined].
Kleothrips (Akleothrips) karimonensis f. parvidens Priesner, 1935c : 334. LECTOTYPE 3, JAvA (SMP),
here designated [examined]. Syn. n.
This is the type-species of Priesner’s subgenus Akleothrips to which he also assigned hardyi and
lacerta. These three and also acanthus comprise the group of species with the median setae on
tergite IX longer than the tube and well-developed anteromarginal setae. The larger males have
bulbous fore femora with an apical tooth and a basal tubercle, also a slim fore tarsal claw. The
smaller males have a smaller basal tubercle on the fore femur, and a longer tube as do the females.
An unspecified number of these small males were named f. parvidens by Priesner. The lectotype
designated here is the specimen labelled as holotype by Priesner.
SPECIMENS STUDIED.
Java: Karimon, holotype 3, paratype 2 of karimonensis, lectotype 3 of f. parvidens, v.1926 (Dammerman
10) (SMF).
Mecynothrips kraussi sp. 0.
(Figs 88, 91, 94, 106, 115)
6 macroptera. Colour dark brown to black, basal stems of antennal segments IJI-V, apices of mid and
hind tibiae, tarsi and cheek spines pale yellow, wings dusky at the base with a dark, median longitudinal
line in the basal half.
Head from behind eyes about 3 times as long as broad, and head prolongation also about 3 times as
long as broad; ocellar setae situated anterior to first ocellus, postocellar setae situated on a pair of tubercles
between the eyes near their posterior margin; 3 to 5 pairs of cheek setae arranged on a cluster of tubercles
immediately behind eyes, 1 pair of cheek setae situated on tubercles in the basal half of head; tentorial
_ pits on ventral surface of head covered by a bilobed tubercle. Antennal segment III 9-7 to 11-4 times as
long as broad and 1-4 times as long as IV. Pronotum slightly longer than broad, anterior angles produced
into upwardly curved horns, anteromarginal and anteroangular setae reduced to minor setae, mid-
laterals small, posteroangular and one pair of epimeral setae well developed. Fore femora enlarged,
rectangular with a dark seta-bearing spine on inner margin at apex, also a tubercle medially and a small
rugose hump at the base; fore tibiae L-shaped with 2 major setae on outer angle, apex slightly produced
inwards under the well-developed, retroussé fore tarsal claw. Median pair of setae on tergite IX about
1-2 times as long as tube. Tube evenly narrowing to apex, about 4-5 times as long as broad, set with a few
small setae.
205
Measurements (holotype 3, with largest and smallest ¢ paratypes, in ym). Body, length 10100 (10945,
9985). Head, length from behind eyes 865 (890, 710); basal breadth 335 (385, 315). Head prolongation,
lateral length 400 (450, 315); basal breadth 125 (135, 125). Anteocellar setae 230/250 (268, 220), post-
ocellar setae 180 (-, —). Postocular setae, pair I 115 (115, 115/135), pair IT 180 (169/190, 155). Antennal
segments, III, length 655 (780, 545); breadth 65 (65, 55); length, IV 480 (545, 415); V 395 (430, -);
VI 240 (270, -); VII 115 (125, -); VIII 95 (115, -). Pronotum, median length 565 (650, 422); maximum
breadth 545 (605, 490); posteroangular setae 120 (160, 110); epimeral setae 142 (170, 135). Fore wing,
length 2975 (3455, 2685); maximum breadth 250 (325, 240); duplicated cilia 59 (67, 52); basal setae, 1,
126 (125, 105); 2, 142 (160, 110); 3, 225 (240, 185). Dorsal setae tergite LX 660 (-, -). Tube, length 545
(615, 545); maximum breadth 125 (135, 125).
2 macroptera. Colour similar to the males.
Head and prolongation not as elongate as the male, only twice as long as broad; ocellar setae situated
lateral or posterior to first ocellus; cheeks set with at most 3 pairs of stout setae in apical half, not ina
cluster, and 1 pair in basal half. Pronotum about 1-4 times as broad as long without horns at anterior
angles; anteromarginal setae small but anteroangulars, mid-laterals, posteroangulars and 2 pairs of epi-
merals well developed. Fore femora not enlarged, unarmed, fore tarsi without a tooth. Tube about 4
times as long as broad.
Measurements (largest and smallest 2 paratypes in um). Body, length 8065, 7485. Head, length from
behind eyes 620, 530; basal breadth 315, 325. Head prolongation, lateral length 240, 190; basal breadth
125, 115. Anteocellar setae 260, —; postocellar setae 190/210, —. Postocular setae, pair I 95/105, 95/115,
pair II 230, —. Antennal segments, III, length 480, —, breadth 45, —; length, IV 375, —; V 305, —; VI 200, -;
VII 95, —; VIII 85, —. Pronotum, median length 385, 325; maximum breadth 500, 480, —; anteroangular
setae 85/95, 40/65; mid-laterals 135, —; posteroangulars —, —; epimerals, outer 115/145, 105; inner —, 85/95.
Fore wing, length 2880, 2305; maximum breadth 250, 190; duplicated cilia 54, 45; basal setae, 1, —, 220;
2, -, -; 3, -; 385. Dorsal setae tergite IX —, —. Tube, length 700, 605; maximum breadth 150, 135.
SPECIMENS STUDIED.
Holotype 3, Solomon Is: Gizo, Titania, 0-20 m, 10.xi.1970 (N. L. H. Krauss) (BMNH).
Paratypes. Solomon Is: 2 3, 2 2, same data as holotype; 3 g, 1 2, same data but 9.xi.1970 (BMNH).
This species is most similar to snodgrassi from which the males may be distinguished by the
arrangement of the cheek setae and the position of the first ocellus. Of this group of species with
median setae on tergite [IX longer than the tube and small pronotal anteromarginal setae, the
only females known are 1 of priesneri, 3 of wallacei and 3 of kraussi. The females of wallacei
have not been examined but those of kraussi may be distinguished from priesneri on the lengths
of the postocular setae and pronotal mid-lateral setae. The median head setae of kraussi are 2:5
times as long as the postoculars, whereas those of priesneri are subequal, and the mid-laterals of
kraussi are less than 1-5 times as long as the anteroangulars, whereas those of priesneri are more
than twice as long.
Mecynothrips lacerta (Priesner) comb. n.
(Fig. 101)
Kleothrips lacerta Priesner, 1935c : 329. LECTOTYPE 2, SumBA (SMF), here designated [examined].
Kleothrips (Synkleothrips) innocens Priesner, 1935c : 331. Holotype ¢, SUMBA (SMF) [examined]. Syn. n.
Kleothrips lacerta was described from a series of female syntypes. The lectotype designated here
is the specimen labelled as holotype by Priesner.
This species belongs to the group with the median setae on tergite IX longer than the tube
and well-developed pronotal anteromarginal setae. K. innocens was described as the type-species
of the subgenus Synkleothrips because of the unarmed fore femora. The holotype is a small male
with a slim fore tarsal claw.
This species is most similar to karimonensis from Java but it may be distinguished mainly
by the more uniform colour of the mid and hind tibiae.
SPECIMENS STUDIED.
Sumba: Kambera, lectotype 2 of /acerta, iii.1925 (Dammerman 6); Laora, 100 m, holotype 3 of innocens,
iv.1925 (Dammerman 89) (SMF).
206
Mecynothrips priesneri Mound
(Figs 96, 104, 105, 107)
Mecynothrips priesneri Mound, 1971b: 281. Holotype 3, NEw GUINEA (BPBM) [1 ¢ paratype examined].
Mecynothrips minor Mound, 19716: 282. Holotype 3, NEw GuINEA (BPBM) [1 3 paratype examined].
Syn. n.
This species belongs to the group with the median pair of setae on tergite IX longer than the tube
and small pronotal anteromarginal setae. Males have large horns on the fore angles of the pro-
notum, rectangular fore femora with an apical tooth and a median tubercle, and a stout fore
tarsal claw. The cover over the tentorial pits on the ventral surface of the head is of one lobe as in
wallacei.
Mound (1971) distinguished the small male, minor, from priesneri by the position of the first
ocellus which is variable, and the major setae on antennal segments III and IV not being at
right angles to the segment. However, these characters are typical of females and small males
throughout the genus.
SPECIMENS STUDIED.
New Guinea: Manus I., Rossur, 150 m, 1 ¢ paratype of priesneri, 30.vi.1959 (J. L. Gressitt); Wau, 1200 m,
1 3, 1 9, 9.xii.1961 (J. H. Sedlacek) (BMNH). New Britain: Gazelle Peninsula, Bainings, St Paul’s,
350 m, 1 3, 7.ix.1955; Kerawat, 60 m, 1 ¢ paratype of minor, 2.xi.1955 (J. L. Gressitt) (BMNH).
Mecynothrips pugilator (Karny) comb. n.
(Fig. 86)
Phoxothrips pugilator Karny, 1913 : 132. Holotype 3, TAIWAN (depositary unknown) [not examined].
Elaphrothrips takahashii Priesner, 1935d: 372. Holotype 2, JAPAN (SMF) [examined]. [Synonymized by
Haga & Okajima 1974 : 379.]
Phoxothrips pugilator Karny; Haga & Okajima, 1974 : 376.
This species belongs to the group with the median setae on tergite IX shorter than the tube. The
large males are distinct in having the inner margin of the fore femur developed into a distinct
bulge. This, however, varies with size and in small males it is indiscernible. Both males and
females of the species may be distinguished from the others in this group, atratus, goliath and
simplex, by the distinctly bicoloured antennae and dark legs. The head is also not so constricted
behind the eyes.
This is the type-species of the monotypic genus Phoxothrips. It was probably described from a
unique male which, according to Haga & Okajima (1974), is not with Karny’s specimens at
Eberswalde. They redescribed the species from a large series of males and females from the
Ryuku Is.
The unique female holotype of takahashii is similar to pugilator although it has a shorter head
prolongation.
SPECIMENS STUDIED.
Japan: Amami Oosima, Gusuku, Loochoo, holotype 2 of takahashii, 27.vii.1932 (S. Minoua) (SMF);
Arakawa, Ishigaki-jima, Rydkyds, 2 d, 1 2 on dead Phoenix (S. Okajima) (BMNH).
Mecynothrips simplex Bagnall comb. rev.
(Figs 90, 99)
Mecynothrips simplex Bagnall, 1912 : 216. Holotype 3, PHILIPPINES (BMNH) [examined].
Kleothrips gigans Schmutz, 1913 : 1058. Syntypes $9, Sri LANKA (Ceylon) [1 ¢ examined]. [Synonymized
by Ananthakrishnan, 1969: 130.]
Dracothrips ceylonicus Bagnall, 1914 : 290. Syntypes 2 3, SRI LANKA (Ceylon) (lost). [Synonymized with
gigans by Bagnall, 1918 : 219.]
Kleothrips agama Priesner, 1935c : 323. Holotype 3, JAVA (SMF) [examined]. [Synonymized by Anantha-
krishnan, 1969 : 130.]
Kleothrips simplex Bagnall; Mound, 1968 : 130.
207
This species belongs to the group in which the median setae on tergite IX are shorter than the
tube. It is most similar to goliath from Sumba except for the slightly paler bases of the anten-
nal segments.
The unique holotype of simplex is a large male with square fore angles to the pronotum but
without horns, and with an enlarged, bulbous fore femur which bears only an apical spine. The
fore tibia has two small teeth at the base and the fore tarsus has a well-developed but slim claw.
The mid and hind tibiae are distinctly bicoloured.
Schmutz (1913) described gigans as the type-species of his genus Kleothrips, differing from
Mecynothrips in lacking horns on the fore angles of the pronotum. The syntype male examined
corresponds to the original illustration in having a large bulbous fore femur with only an apical
tooth, a tooth at the base of the fore tibia and a well-developed slim fore tarsal claw. It is in these
characteristics inseparable from simplex.
The specific synonymy of ceylonicus with gigans was not stated but implied. In March 1918
Bagnall synonymized his monotypic genus Dracothrips with the, at that time, monotypic Kleo-
thrips. There were no other species in Kleothrips until Hood described acanthus in October 1918.
Priesner (1935) described agama from a large series of males and females from Java. They
differ from simplex only in the lack of any small teeth at the base of the fore tibia. From the
material examined this character would appear to vary according to the stoutness of the specimen.
Ananthakrishnan (1969) discussed and illustrated the structural variation in the head and fore
legs of this species, but unfortunately used the name gigans rather than the older name simplex.
SPECIMENS STUDIED.
Sri Lanka (Ceylon): 1 3 (E. E. Green); 1 3, 8.iv.1907 (B. Fletcher); 1 3 (Thwaites); Peradeniya, 1 2 (E. E.
Green), 1 3 on dead Hevea leaves, 10.v.1914; ? Ceylon: 1 ¢ (BMNH). India: Adichanallur, 1 3, 1 2 on
Areca sheath, 25.vi.1967 (TNA); Uyithri, 2 3, 4 2 on dry twig, 15.viii.1969 (TNA) (BMNH). Malaya:
Penang, 1 92, 20.xi.1945 (H. T. Pagden) (BMNH). Java: Karimon, holotype 3g, paratype 2 (+2 3 paratypes,
2 2 BMNH) v.1926 (Dammerman 10) (SMF); Bogor, 2 3, 2 2 on dead leaves, 25.x.1973 (L. A. Mound);
Jasinga Forest Reserve, 2 gj on dead leafy branch, 30.x.1973 (L. A. Mound) (BMNH). Borneo: W. Sarawak,
Quop, | J, iii.1914 (G. E. Bryant) (BMNH). Philippines: holotype ¢ of simplex (E. Simon) (BMNH).
Mecynothrips snodgrassi Hood
(Fig. 92)
Mecynothrips snodgrassi Hood, 1952 : 294. Holotype 3, SOLOMON Is (USNM) [not examined].
This species belongs to the group which has the median setae on tergite IX longer than the tube,
and small pronotal anteromarginal setae. The males have large horns at the fore angles of the
pronotum, and rectangular rather than bulbous fore femora with an apical tooth and a median
tubercle on the inner margin. The fore tarsal claw is well developed and there is a bilobed cover
over the tentorial pits on the ventral surface of the head. The original illustration of the large
holotype male shows the irregular arrangement of cheek spines and the bilobed process on the
head. In these characteristics this species closely resembles the new species Kraussi, also from the
Solomon Is, but it may be distinguished by the more numerous cheek setae in the basal half,
kraussi having only 1 pair, and the stouter fore tarsal claw. The position of the first ocellus,
although variable, is apparently anterior to the ocellar setae in snodgrassi and usually posterior in
kraussi.
SPECIMEN STUDIED.
Solomon Is: Florida Group, Tulagi, 1 3, 2.ix.1960 (C. W. O’Brien) (BMNH).
Mecynothrips wallacei Bagnall
(Fig. 95)
Mecynothrips wallacei Bagnall, 1908a : 357. Holotype 3, NEw GUINEA (BMNH) [examined].
Mecynothrips magnus Girault, 1929b: 1. Syntypes 3, AUSTRALIA (QM) [not examined]. [Synonymized
by Mound, 1971 : 283.]
208
Mecynothrips bagnalli Priesner, 1935c : 335. Holotype 9, Ker Is (? SMF) [not examined]. [Synonymized
by Mound, 1971 : 283.]
Mecynothrips bagnalli f. imbecilla Priesner, 1935c : 338. Syntypes g2, Ker Is (? SMF) [not examined].
Syn. n.
This is the type-species of the genus Mecynothrips. It belongs to the group of species with small
anteromarginal prothoracic setae and the median setae on tergite IX longer than the tube.
The holotype of wallacei is a large male with well-developed pronotal horns, rectangular fore
femora with an apical spine and a tubercle on the inner margin just in the apical half. The fore
tibiae have a slight bulge at the base and are drawn out into a tubercle at the apex. The fore
tarsal claw is stout and well developed. The fore wings are pale apart from a dark horizontal
median band in the basal half. There is a cover over the tentorial pits on the ventral surface of
the head which is of one lobe, as in priesneri.
Priesner described the form imbecilla from small males which have shorter pronotal horns
as well as smaller fore femoral spines and a smaller fore tibial apical tubercle. There is a small
male possibly of wallacei in the BMNH from New Guinea which has a small slim fore tarsal
claw and only an apical spine on the fore femur. The female which was apparently collected with
this male, however, has well-developed anteromarginal prothoracic setae.
SPECIMENS STUDIED.
West Malaysia (‘Malayischen Archipel.’): 1 ¢ (? Karny) (BMNH). New Guinea: Dorey, holotype 3 of
wallacei (Wallace); Port Moresby, Waigani, 1 3, 1 9, leaf litter, 28.ix.1969 (BMNH).
MEIOTHRIPS Priesner
Idolothrips (Meiothrips) Priesner, 1929b: 197. Type-species: Idolothrips (Meiothrips) annulatus Priesner
(here regarded as a synonym of Acanthinothrips annulipes Bagnall), by monotypy.
Meiothrips Priesner; Bagnall, 1934a : 494.
Meiothrips (Telothrips) Kudo & Ananthakrishnan, 1974: 385. Type-species: Meiothrips (Telothrips)
nepalensis Kudo & Ananthakrishnan, by monotypy. [Junior homonym of Telothrips Priesner, 1929a.]
Syn. n.
Meiothrips (Aculeathrips) Kudo, 1975 : 421. Replacement name for Telothrips Kudo & Ananthakrishnan.
Syn. n.
Large, elongate species of Idolothripini. Head long, projecting in front of eyes; first ocellus between bases
of antennae; inter- and postocellar setae well developed; vertex with 1 pair of postocular setae close
together, well behind eyes; cheeks with at least 2 pairs of short stout setae. Antennae 8-segmented,
exceptionally elongate, III as long as fore tibia; III with 2 sense cones, IV with 4 sense cones. Pronotum
with epimeral sutures not complete; praepectal plates weakly developed, lateral to mouth cone. Fore
tarsi unarmed; femora slender but irregularly swollen distally, bearing 4 pairs of stout capitate setae.
Mesonotal lateral setae, metanotal median setae and metepimeral setae stout and elongate. Fore wing
with or without duplicated cilia; sub-basal setae relatively small, II shortest, III longest. Pelta pointed
medially, with broad lateral wings. Tergites III-VI with antecostal ridge recurved medially; tergites with
2 pairs of sigmoid wing-retaining setae and several accessory setae laterally, one or more pairs of these
setae sometimes on or anterior to antecostal ridge; setae on IX short; tube with numerous erect setae,
with or without tubercles or denticles in 3.
This genus is closely related to Bactridothrips and Idolothrips. In all three genera the species have
numerous erect setae on the tube, and the metanotum and metepimera bear a pair of stout setae.
The females of Meiothrips species can be distinguished from females of species in the other two
_ genera mainly by the longer antennae and more slender head. The males of M. annulipes, like
the males of Idolothrips species, have the posteroangular setae of the abdominal tergites stout and
thorn-like. Moreover, in both M. nepalensis and I. dissimilis the dorsal surface of the tube bears
two rows of stout tubercles, whereas in M. annulipes the dorsal surface of the tube bears numerous
small denticles. The subgenus Aculeathrips is thus intermediate between Meiothrips and Idolo-
thrips, and it is possible that only one genus is needed for this whole group of species. The species
of Meiothrips have been collected infrequently, but the genus is widespread between Nepal,
southern India and Borneo. In contrast Idolothrips is still known only from Australia.
209
Key to species
1 Fore wing without duplicated cilia; pelta with numerous dentate microtrichia on lines of
sculpture in posterior half (Fig. 111); mid-femora uniformly dark brown; ¢ with 1 pair of
posteroangular setae short and thorn-like on tergites VII-VIII; tube of ¢ with numerous
small denticles on dorsal surface [Malaya, Sumatra, Borneo] . ; ; annulipes (p. 210)
— Fore wing with at least 12 duplicated cilia; sculpture on pelta not bearing microtrichia; mid-
femora yellow in part; 3 without thorn-like tergal posteroangular setae . A Z
2 Tube of 3 with 2 longitudinal rows of pointed tubercles dorsally; setae B, on tergite IX of 3
shorter than width of tergite; antennal segments VI-VII with a short apical, ventral pro-
longation (Fig. 110); femoral major setae yellow [Nepal, Thailand]. ‘ nepalensis (p. 212)
— Tube of 3 without dorsal tubercles or denticles; setae B, on tergite IX of 3 longer than width
of tergite; antennal segments VI-VII not prolonged ventrally (Fig. 109); femoral major
setae brown to dark brown [India, Thailand, Malaya] . : ; : . menoni (p. 210)
Meiothrips annulipes (Bagnall)
(Figs 108, 111-112)
Acanthinothrips annulipes Bagnall, 1914 : 378-379. Lectotype 3, SARAWAK (BMNH) [examined].
Idolothrips (Meiothrips) annulatus Priesner, 1929b : 197-201. Syntypes 93, SUMATRA (SMF) [1 ¢ BMNH
examined]. Syn. n.
Meiothrips annulipes (Bagnall) Bagnall, 1934a : 494.
The specimens in the type-series of annulipes are larger and more brightly coloured than the small
teneral male syntype of annulatus which has been studied. The degree of development of the thorn-
like setae on the posterior angles of the tergites is variable as in Jdolothrips (Mound, 1974).
Priesner pointed out in the original description of annulatus that on tergite V it is not unusual
for the seta on one side to be long and ‘female-like’, but on the other side to be short and stout.
This, however, is not true of larger males and it can be inferred that on smaller males both setae
on tergite V will be slender.
SPECIMENS STUDIED.
Borneo: Sarawak, Mt Matang, lectotype 3 of annulipes on dead wood at 1000 ft (30 m], 13.xii.1913
(G. E. Bryant) (BMNH); 6 3, 1 2 paralectotypes with similar data, xii.1913-i.1914 (BMNH). Sumatra:
Mentawei, Siberat, syntype 3 of annulatus, 10.ix.1924 (Karny, 25) (BMNH). Malaya: Tapah, 1 3, 1 2 on
dead leaves, 30.vii.1976 (Okajima) (SO, Tokyo).
Meiothrips menoni Ananthakrishnan
(Fig. 109)
Meiothrips menoni Ananthakrishnan, 1964 : 99-101. Holotype 9, InpIA (TNA) [not examined].
This species was described from a single female collected at Trichur in southern India. That
specimen and the three specimens from India listed below have the tibiae largely yellow except
for a pale brown mark near the base and an even fainter mark near the apex. In contrast, the
specimens listed below from Malaya have an extensive dark brown sub-basal area on the tibiae,
and a narrow dark brown area subapically. Moreover, these specimens have short, dark pronotal
posteroangular and anteromarginal setae, and the setae on the vertex and ocellar region are
also short and dark, whereas the Indian specimens have longer and paler setae. If only these
two samples were available then the Malayan specimens would undoubtedly be regarded as a
distinct species. However, the two specimens listed below from Thailand have the tibiae inter-
mediate in colour between the Indian and Malayan specimens, and moreover the pronotal setae
are long and pale whereas the head setae are short and dark. These specimens suggest that
menoni exists as a cline between southern india and Malaya; therefore no attempt is made here to
distinguish between these populations by the use of formal names.
SPECIMENS STUDIED.
India: Trichur [Kerala], 1 3, beating logs, 7.xi.1963; Kallar [? Mysore], 1 3, 1 2 on Areca leaf, 9.ix.1967
210
108-112, Meiothrips species. (108) M. annulipes. 109, 110, antennal segments VI-VIII,
Figs 108-115
(109) M. menoni; (110) M. nepalensis. 111-112, M. annulipes, (111) pelta; (112) tergite V. 113-115,
Mecynothrips species pronota, (113) M. hardyi; (114) M. atratus (zuluensis holotype); (115) M.
kraussi.
211
(TNA). Thailand: Doi Suthep, 1 3, 1 2 from dead leaves, 1100 m, 11.viii.1976 (Okajima) (SO, Tokyo).
Malaya: Tapah, | 3 from dead leaves, 28.vii.1976 (Okajima) (SO, Tokyo); Kuala Lumpur, 13 2, 7 ¢ from
dead branches, 23-26.xii.1969 (Andre) (BMNH).
Meiothrips nepalensis Kudo & Ananthakrishnan
(Fig. 110)
Meiothrips (Telothrips) nepalensis Kudo & Ananthakrishnan, 1974 : 385-387. Syntypes 39, NEPAL (2 3
syntypes BMNH) [examined].
Meiothrips (Aculeathrips) nepalensis Kudo & Ananthakrishnan; Kudo, 1975: 421.
As discussed above, the remarkable dentate structure of the tube in this species is not unique but
also occurs in Idolothrips dissimilis Girault from Australia. One of the males listed below from
Thailand is exceptionally small. This specimen differs from the other nepalensis males in having
no long metanotal setae, and in having few small denticles near the base of the tube although the
dorsal tubercles are well developed. It is not possible to know if this is normal variation, aberration
or another new species. The pronotal anteromarginal setae are shorter than the mid-lateral setae
in one of the four syntypes which have been studied, but this is not so in the other three syntypes.
SPECIMENS STUDIED.
Nepal: Dhunche, 4 3 syntypes on dry leaf, 19.x.1973 (Kudo) (2, BMNH). Thailand: Doi Suthep, Chieng-
mai, 4 9, 2 on dead leaves, 22.iv.1976 (Suzuki) (BMNH); Doi Suthep, 1100 m, 1 3, 11.viii.1976 (Okajima)
(SO, Tokyo).
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Haga, K. 1972. Thysanoptera occurring on dead leaves and dead branches. Rept high sch. Osaka Kyoika
Univ. 14: 100-113.
Haga, K. & Okajima, S. 1974. Redescription and status of the genus Phoxothrips Karny (Thysanoptera).
Kontyii 42 : 375-384.
Heeger, E. 1852. Beitriige zur Naturgeschichte der Physapoden (Blasenfusse). Sher. Akad. Wiss. Wien
9: 123-141.
Hood, J. D. 1918. A new Kleothrips (Thysanoptera) from North Queensland. Bull. Brooklyn ent. Soc.
13 : 77-79.
— 1919. On some new Idolothripidae. Insecutor Inscit. menstr. 7 : 66-74.
— 1927. A contribution toward the knowledge of New York Thysanoptera, with descriptions of new
genera and species. Entomologica am. 7: 209-245.
— 1935. A note on heterogony in the Thysanoptera, with descriptions of two new species from Tan-
ganyika. Stylops 4: 193-201.
— 1952. Mecynothrips snodgrassi, a new thrips from the Solomon Islands. Proc. ent, Soc. Wash.
54: 294-301.
Ishida, M. 1932. Fauna of the Thysanoptera in Japan. Insecta matsum. 7: 1-16.
— 1934. Fauna of the Thysanoptera in Japan. Insecta matsum. 8: 147-151.
— 1936. Fauna of the Thysanoptera in Japan. Insecta matsum. 10: 154-159.
Jacot-Guillarmod, C. 1939. New species of Phlaeothripidae (Thysanoptera) from South Africa. J. ent.
Soc. sth. Afr. 1: 47-77.
Karny, H. 1912. Ueber einige afrikanische Thysanopteren. Fauna exot. 2 (5): 19-20, 22-26.
— 1913. H. Sauter’s Formosa Ausbeute Thysanoptera. Supplta ent. 2: 127-134.
— 1920a. Die exotischen Tubiliferen (Thysanoptera) des Deutschen Entomologischen Museums
(Berlin-Dahlem). Ent. Mitt. 9: 88-94, 104-111, 186-191.
— 1920b. Some Thysanoptera from the Philippine Islands. Philipp. J. Sci. 17: 203-209.
— 1920c. Nova Australska Thysanoptera, jes nasbiral Mjéberg. Cas. ésl. Spol. ent. 17 : 35-44.
—— 1921a. Beitrige zur Malayischen Thysanopterenfauna I-III. Treubia 1: 277-291.
—— 1921b. Beitrage zur Malayischen Thysanopterenfauna IV-V. Treubia 2 : 37-83.
—— 1923. Beitrage zur Malayischen Thysanopterenfauna VI-VIII. Treubia 3 : 277-380.
— 1924. Results of Dr. E. Mjéberg’s Swedish Scientific Expeditions to Australia 1910-1913. Thysano-
ptera. Ark. Zool. 17 (A) 2: 1-56.
— 1925. On some tropical Thysanoptera. Bull. ent. Res. 16: 125-142.
Kudo, I. 1975. New name for the subgenus Te/othrips Kudo et Ananthakrishnan, not Priesner (Thysano-
ptera, Phlaeothripidae). Kontyii 43: 421.
Kudo, I. & Ananthakrishnan, T. N. 1974. A new subgenus and species of Meiothrips Priesner (Thysano-
ptera, Megathripinae) from Nepal. Kontyé 42 : 385-387.
Kurosawa, M. 1932. Descriptions of three new thrips from Japan. Kontyé 5 : 230-242.
Moulton, D. 1928a. New Thysanoptera from Formosa. Trans. nat. Hist. Soc. Formosa 18: 287-328.
7a ke
—— 1928b. New Thysanoptera from India. Indian Forest Rec. 13 : 285-292.
—— 1933. New Thysanoptera from India. Indian Forest Rec. 19: 1-6.
—— 1947. Thysanoptera from New Guinea, the Philippine Islands and the Malay Peninsula. Pan-Pacif.
Ent. 23: 172-180.
Mound. L. A. 1968. A review of R. S. Bagnall’s Thysanoptera Collections. Bull. Br. Mus. nat. Hist. (Ent.)
Suppl. 11: 1-181.
1970. Thysanoptera from the Solomon Islands. Bull. Br. Mus. nat. Hist. (Ent.) 24: 83-126.
—— 197la. Gall-forming thrips and allied species (Thysanoptera: Phlaeothripinae) from Acacia trees
in Australia. Bull. Br. Mus. nat. Hist. (Ent.) 25 : 387-466.
—— 1971b. A review of the Melanesian genus Mecynothrips (Thysanoptera). J. nat. Hist. 5: 279-283.
—— 1974. Spore feeding thrips (Phlaeothripidae) from leaf litter and dead wood in Australia. Aust. J.
Zool. Suppl. 27 : 1-106.
Pergande. 1896. Description of a new species of Jdolothrips. Ent. News 7: 63-64.
Priesner, H. 1928a. Thysanopterologica III. Zool. Jb. 56 : 43-66.
— 19285. Uber australische Thysanopteren. Sber. Akad. Wiss. Wien Abt. 1 137: 643-659.
—— 1929a. Bernstein Thysanoptera II. Bernstein Forsch. 1: 111-138.
— 1929b. Spolia Mentawiensia: Thysanoptera. Treubia 11: 187-210.
— 1930. Indomalayische Thysanoptera II. Treubia 11 : 357-371.
—— 1932. Thysanopteren aus dem Belgischen Congo. Revue Zool. Bot. afr. 22 : 320-344.
—— 1934a. Indomalayische Thysanopteren VI (I-I]). Konowia 13 : 193-202.
—— 19346. Indomalayische Thysanopteren VI (IID). Konowia 14: 58-67.
—— 1935a. Indomalayischen Thysanopteren VI (III Fortsetzung). Konowia 14: 159-174.
—— 1935b. Indomalayischen Thysanopteren VI (III Fortsetzung). Konowia 14: 241-255.
— 1935c. Indomalayischen Thysanopteren VI (III Schluss). Konowia 14 : 323-339.
— 1935d. New or little-known Oriental Thysanoptera. Philipp. J. Sci. 57: 351-375.
—— 1939. Thysanopterologica (VIII). Proc. R. ent. Soc. Lond. (B) 8 : 73-78.
—— 1940. Thysanopterologica IX. Philipp. J. Sci. 71: 403-411.
—— 1959. On the genus Dinothrips Bgn. Idea 12 : 52-59.
Ramakrishna Ayyar, T. V. 1930. Notes on Indian Thysanoptera, with descriptions of new species. Rec.
Indian Mus. 36 : 491-498.
1934. Entomological investigations on the spike disease of Sandal (21), Thysanoptera. Indian
Forest Rec. 20 (4): 1-12.
Schmutz, K. 1913. Zur Kentniss der Thysanopterenfauna von Ceylon. Sber. Akad. Wiss. Wien 122: 991-
1089.
Seshadri, A. R. & Ananthakrishnan, T. N. 1954. Some new Indian Thysanoptera I. Indian J. Ent. 16 : 210-
256.
Takahashi, R. 1936. Thysanoptera of Formosa. Philipp. J. Sci. 60 : 427-459.
—— 1937. Descriptions of new Thysanoptera from Formosa, with notes on the species found on the
high elevations of the island. Tenthredo 1 : 339-350.
Vuillet, A. 1914. Description d’un Dicaiothrips nouveau de l’Inde (Thysanopt. Idolothripidae). Bull. Soc.
ent. Fr. 1914 : 276-278.
Woo, K. S. 1974. Thysanoptera of Korea. Korean J. Ent. 4: 1-90.
Index
Synonyms are in italics
acanthus, 202
achaetus, 183
Acrothrips, 199
Aculeathrips, 209
Adiaphorothrips, 186
Aesthesiothrips, 163
affinis, 169
agama, 207
agasthya, 172
Akainothrips, 186
Akleothrips, 199
amoenus, 176
andrei, 158
annulatus, 210
annulipes, 210
anodon, 169
antennatus, 189
apertus, 172
approximatus, 183
artocarpi, 189
athletes, 176
Atractothrips, 196
214
atratus, 203
bagnalli, Ecacanthothrips, 161
bagnalli, Mecynothrips, 209
bakeri, 176
beesoni, 179
bicolorisetosus, 190
biuncinatus, 190
bouvieri, 180
braueri, 190
bruneitarsis, 182
bryanti, 161
buffai, 193
celebensis, 169
celosia, 192
ceylonicus, 207
chandana, 183
clarispinis, 184
clavipes, 164
Cnemidothrips, 163
Cnestrothrips, 186
coniger, 160
consanguineus, 162
coreanus, 178
coronatus, 182
corticosus, 192
coxalis, 161
Cradothrips, 171
crassiceps, Ecacanthothrips,
161
crassiceps, Dicaiothrips, 169,
172
curvipes, 178
dallatorensis, 183
dammermani, 189
decorus, 193
denticollis, 179
depokensis, 178
diabolus, 193
Diaphorothrips, 163
Dicaiothrips, 171
Dinothrips, 166
Dracothrips, 199
Ecacanthothrips, 156
Elaphoxothrips, 171
Elaphridia, 172
Elaphrothrips, 171
eranthemi, 183
erythrinus, 162
flavipes, 161
fletcheri, 162
formosensis, 162
fulmeki, Dinothrips, 170
fulmeki, Elaphrothrips, 180
Surcifer, 170
gardneri, 169
gargantua, 202
gigans, 207
giganteus, 203
goliath, 203
graveleyi, 184
greeni, 180
guamensis, 196
guineaensis, 161
hamipes, 164
haplodon, 193
hardyi, 203
heveae, 194
Hoplandrothrips, 156
Horistothrips, 174
hystrix, 182
imbecilla, 209
imitator, 178
indicus, 194
inermis, 161
innocens, 206
insignis, 181
insularis, 181
io, 162
isshikii, 186
jacobsoni, Dinothrips, 169
jacobsoni, Elaphrothrips, 181
jatrophae, 163
juglandis, 167
kanoi, 205
karimonensis, 205
karnyi, Elaphrothrips, 178
karnyi, Machatothrips, 192
kemneri, 169
Kleothrips, 199
kraussi, 205
lacerta, 206
leai, 160
lentus, 194
levis, 182
longicauda, 167
Machatothrips, 186
Macrothrips, 186
magnus, 208
mahensis, 182
malayensis, Elaphrothrips, 182
malayensis, Malesiathrips, 198
Malesiathrips, 196
malloti, 169
matsumurai, 162
Mecynothrips, 199
Meiothrips, 209
menoni, 210
mentaweiensis, 176
micidus, 180
minor, 207
monodon, 167
montanus, 190
mucronatus, 179
multidens, 190
215
nepalensis, 212
notabilis, 182
novus, 183
obscuricornis, 179
Ormothrips, 156
pallicornis, 186
parvidens, 205
paucidens, 190
Paxillothrips, 166
Dhilippinensis, 162
Phoxothrips, 199
priesneri, Ecacanthothrips, 162
priesneri, Mecynothrips, 207
procer, 183
productus, 179
proximus, 183
pugilator, 207
quadrudentatus, 195
ramakrishnai, 162
rex, 182
sanguineus, 161
secus, 178
sensitivus, 183
serex, 202
seychellensis, 184
silvaticus, 195
simplex, Machatothrips, 190
simplex, Mecynothrips, 207
simplicidens, 193
snodgrassi, 208
solomoni, 199
sorex, 199
spatiata, 194
spiniceps, 184
spinipes, 161
spinosus, Diaphorothrips, 164
spinosus, Dinothrips, 169
Steinskyi, 161
sumatrensis, 170
sumbanus, 179
Synkleothrips, 199
takahashii, 207
Telothrips, 209
thevetii, 164
thoreauini, 174
tibialis, 161
unguipes, 164
wallacei, 208
zuluensis, 203
5
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: Bulletin of the
British Museum (Natural History)
The phlebotomine sandflies (Diptera:
Psychodidae) of the Oriental Region
D. J. Lewis
Entomology series Vol 37 No 6 30 November 1978
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This number completes Volume 37
ISSN 0524-6431 Entomology series
Vol 37 No 6 pp 217-343
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 30 November 1978
The phlebotomine sandflies (Diptera : Psychodidae)
of the Oriental Region
D. J. Lewis |
Department of Entomology, British Museum (Natural History), Cromwell Road,
London SW7 5BD
Synopsis
Introduction .
Explanation of terms
Methods ;
Taxonomic features
Classification
Key to the species and subspecies of Oriental Phlebotominae
Taxonomy and distribution of species
Contents
Genus Phlebotomus Rondani & Berté
Subgenus Phlebotomus Rondani & Berté
Subgenus Paraphlebotomus Theodor .
Subgenus Synphlebotomus Theodor
Subgenus Larroussius Nitzulescu
Subgenus, Adlerius Nitzulescu
Subgenus Euphlebotomus Theodor
Subgenus Anaphlebotomus Theodor
Ungrouped
Subgenus Idiophlebotomus ‘Quate & Fairchild
Genus Sergentomyia Franca & Parrot
The musai-group
Subgenus Sergentomyia Franca & Parrot
Subgenus Parrotomyia Theodor
Subgenus Grassomyia Theodor
Subgenus Neophlebotomus Franca & Parrot
The nicnic-group
Ungrouped :
Subgenus Sintonius Nitzulescu ,
Nomen nudum
Aspects of biology
General distribution
Breeding habits .
Adult resting sites
Food of adults
Seasonal prevalence
Natural enemies
Relation to disease
The effect of bites
Viruses :
Dermal leishmaniasis ;
Visceral leishmaniasis
Acknowledgements
References
Index
Bull. Br. Mus. nat. Hist. (Ent.) 37(6) : 217-343
217
218
218
219
219
22)
Zee
223
232
233
Pas 8)
235
236
237
239
240
247
250
250
252
253
223
256
268
269
294
295
307
311
311
311
318
318
319
321
aed
S22
a Pw A
oan
322
323
331
331
341
Issued 30 November 1978
Synopsis
A key is given for the 122 species and two subspecies of Phlebotominae known from the Oriental Region,
which are grouped in two genera. Taxonomic citations include all those necessary for reference to early
works. Seventeen new species and one subspecies are described and named, and three species are de-
scribed and given vernacular names. Many existing descriptions are amplified.
Detailed study showed the significance of some structures not normally used in taxonomy, including
features of the stylets and sculpture of the pigment patch. Long-standing problems of Phlebotomus
argentipes, the Sergentomyia babu complex, S. malabarica and S. perturbans were investigated. Variation
in P. argentipes is associated with biological differences in host choice and disease transmission. Many new
localities are recorded and the distribution of each species is mapped. Aspects of biology are reviewed
with special reference to general distribution, biting habits and seasonal distribution. Medical importance
is summarized.
Introduction
Only three species of Phlebotominae were known in the world in 1905 when impetus was given
to their study by their suspected association with sandfly fever and leishmaniasis. By 1925 47
more species were known, 14 of them from the Orient, including nine from India and two from
Sri Lanka. In 1926 Adler and Theodor discovered the taxonomic value of cibarial teeth and
spermathecae, giving further impetus to sandfly taxonomy, and by 1970 649 taxonomic names
had been proposed. Devastating epidemics of kala-azar, discussed in a later section, had led to
intensive research during which J. A. Sinton published a long series of important taxonomic
papers, mainly on India and Pakistan, from 1921 to 1933. Raynal and Gaschen investigated
Indo-Chinese species, a few other workers studied some Oriental species, and Quate recently
made important contributions from south-east Asia. Otherwise, interest waned because kala-azar
came under control, and was not present in south-east Asia to stimulate research. Recently,
however, several factors have rekindled interest in Oriental Phlebotominae and necessitated a
taxonomic review. Reduction in anti-malarial house-spraying has led to some recrudescence of
kala-azar. Several viruses have been found in sandflies, and the modern interest in zoonoses,
animal reservoirs of leishmaniasis, and possible additional vectors has led to much collecting in
outdoor habitats. Improved trapping methods have yielded large collections for study.
In the present work 17 species and one subspecies are described as new, some by D. J. L. &
J. Jeffery and the others by D. J. L. Several existing descriptions are amplified and the males of
some species are described for the first time. The 124 taxa now known from the Region comprise
122 species and two subspecies. More species may exist, for Quate & Fairchild (1961) expected
more to be found in West (Peninsular) Malaysia and Borneo, and Quate (1965) trebled the number
of known Philippine species, and estimated that a hundred might exist.
In the section on biology several aspects are summarized, including seasonal occurrence which
is important for planning future surveys.
The data of specimens critically examined for the descriptions and measurements, including
the type-specimens of new species and subspecies, are listed under ‘Material examined’; all such
specimens are in the BMNH unless otherwise stated. The data of type-specimens of previously
described species are given in the synonymies and not repeated under ‘Material examined’.
Under the heading ‘Distribution’ are listed data from several sources which can be dis-
tinguished by the form of annotation. Localities of specimens routinely identified for distribution
records (but not used for the descriptions and measurements) are indicated by ‘(BMNH)’ or by
the collector’s name in parentheses; in the latter case at least some of the specimens seen are also
in the BMNH unless otherwise indicated. Published records are indicated by a reference to the
published source. Records taken from the manuscript notes of J. A. Sinton are indicated by
‘(Sinton’s notes)’. Localities of specimens already listed under ‘Material examined’ are marked
‘(as above)’. Dates of collection and other data are added only when thought to be of particular
interest.
The length of the scale lines on the figures is given in millimetres.
218
Explanation of terms
Abbreviations and some terms are explained below
Antenna 3 etc. Antennal segment 3 etc.
BMNH British Museum (Natural History)
BPBM Bernice P. Bishop Museum, Honolulu
Complex A group of closely similar species or infraspecific forms
ICZN International Code of Zoological Nomenclature
R, etc. Wing veins: radius branch 2 etc.
Series A group of species within a subgenus or species-group
Species-group A group equivalent to a subgenus
TC Theodor collection, Hadassah-Hebrew University Medical School, Jerusalem
(location of types shown by personal communication, 1977)
ZMA Zoological Museum, Amsterdam University
Methods
Many mounting media have been used for sandflies, and for the present work a variant of
Berlese’s medium (Lewis, 19735) was employed which can be filtered through previously wetted
filter paper during preparation. The medium is useful for rapid processing of large numbers of
flies. To allow for shrinkage through evaporation, either specimens were left for some hours before
addition of a cover glass, or mounts were topped up with mounting fluid after a few hours or a
day. Slides were kept at about 37 °C, usually for about five weeks, till the peripheral mountant
was brittle and no movement in thick mounts was noticed, but not long enough for cover glasses
to become loose. Excess mountant was then removed with a needle, and cover glasses were
ringed with Euparal which is more secure than the commonly used nail varnish.
Drying alters the refractive index and reduces the initial clarity of some ascoids and sperma-
thecae, but this can be restored by re-wetting. If mounts are insufficiently dried Euparal may mix
with the mountant, so that preparations become grey and specimens fragile. This process can
take 20 years or more; it is a warning against adopting new methods without careful consideration.
Caustic potash (10 per cent) was not used for maceration, but the head of one old pinned fly
was put in hot potash for one minute because dried tissues had distorted the cibarium.
It is often necessary to remount a sandfly, for reorientation, for examining spermathecae, or
for studying such structures as the cibarial teeth of S. cheongi and the lower parameral lobes of
P. philippinensis. For recovering a specimen from a sealed mount, a slide was briefly immersed in
water, and Euparal was chipped away. A small square of cover glass above the specimen was
isolated by thrusting a needle, plough-like, beneath the glass, and the water and debris were
washed away and replaced by Berlese’s medium. A fresh cover glass was added to keep the mount
wet till it softened in a few hours or over-night. Water could be dropped on, or pipetted in, for
rapid soaking, but might form an inconvenient precipitate or wash away small objects. Mounts,
used thus for recovering specimens, serve as storage capsules which are better than vials of
alcohol, for space-saving, upkeep, retrieval and inspection.
When sandflies in a collection are being mounted time can be saved if one or a group of species
can be isolated provisionally from the others by means of external features. This can occasionally
be done by observing size (large, medium or small), labrum length, dark or pale scutum or abdo-
men, and broad or narrow wings. Species provisionally separable in this way include S. gombaki,
S. iyengari and S. malayae.
Identification of sandflies is often difficult for several reasons. For example, the appearance of
internal taxonomic features may be affected by the condition of a fly, method of mounting, or
degree of illumination. It is often difficult to associate the sexes, and the problem of finding larvae
limits their use.
Measurements were made from Berlese mounts and did not alter appreciably with drying.
Wing measurements could be slightly different if flies were examined in water.
If eyes were unusually large or small, their length, parallel to the head axis, was measured and
divided by the head length measured from the clypeus tip to the level of the postero-lateral
margins.
219
The cibarial teeth were counted in heads not subject to pressure. Compression may reveal a
few more lateral teeth or spicules but spoils a specimen. Cibarial teeth numbering more than 60,
often seen in subgenus Parrotomyia, are difficult to count unless they are magnified about 300
times and ticked on a camera lucida image. In some species with about 90 teeth the width of one
was estimated by examining the median ones at about x 2000 and counting the number in a
length of 0-008 mm.
If the inter-arcal area is dark or refractive it is occasionally necessary to separate the pigment
patch from the ventral wall of the cibarium. This can be done by mounting the relevant part of
the cibarium upside down in Berlese’s medium and sliding the armature forward.
It is sometimes useful to examine a cibarium in longitudinal or transverse optical section. For
this purpose a head, with antennae removed to facilitate orientation, may be left over-night in a
drop of Berlese’s medium, in which the soft parts become invisible but remain to preserve the
shape. The drop is then wetted with fresh medium to prevent shearing, and the head cut in two
with a razor blade. This, held with both hands, can be directed to within 0-02 mm or less of the
objective. For lateral viewing the eyes are then removed for clarity and to facilitate orientation.
Preparation of the stylets of some species needs special care. For example, the maxillary
lacinia of S. iyengari and the tip of a hypopharynx or a mandible are almost indistinguishable
under a dissecting microscope, and are best mounted in situ on a slide after removal of other
structures. Fortunately these time-consuming procedures are seldom necessary in routine work.
Much detail of the fascicle can be seen if the labium is removed by drawing it sideways and back-
wards off the head.
Some antennal ascoids are very delicate, with their tips touching the antenna, and some distal
ascoids can be small. In such cases the antennae are best mounted temporarily in water. The small
ascoids of S. malayae are difficult to expose but can be seen well in occasional specimens. One
ascoid was measured for each species, without the spur if present.
The presence or absence of a papilla on antenna 5S is of interest in some species. Its preparation
and inspection sometimes take time if a fly is delicate or has been macerated in potash and
mounted in balsam. In such a case papillae may be invisible, even under phase contrast, and must
be remounted.
Palp segments are unsuitable for measuring if bent or shrunk. In this case a head may be
temporarily mounted top-upward in dilute Berlese’s medium so that the palps lie flat and expand.
If spermathecae are not visible in whole mounts the tip of the abdomen (sixth and succeeding
segments) may be detached and slightly compressed from above (with a backward and forward
movement) to release eggs or other loose objects and spread the segments. It is sometimes tempt-
ing to clear with potash but one should remember that the spermatheca used by the taxonomist
is but a small and often delicate part of the whole spermathecal tissue (Theodor, 1965 : figs 3,
5E) and can easily be distorted by removal of soft parts. If ducts are delicate they can sometimes
be exposed by removing virtually all external sclerites except the bifid sternum 8.
Complete descriptions of taxa would be too long. It is often necessary to omit mention of
some group characters, absence of features which occur only in a few species, some features
shown by figures, and some structures too small for regular inspection or only seen properly
with the electron microscope. No description of a single species, therefore, can serve as a model,
but the following list of structures includes most of those used in the present work.
9. Head: labrum length and its relation to wing length, labral brush, labral apical, subapical, adoral and
cibarial sensilla; cibarial shape and pigmentation, armature, pigment patch and arch, and relative width
of the underlying salivary pump; pharyngeal shape and armature of the dorsal, and sometimes ventral,
plates; hypopharyngeal teeth; antenna 3 (length and its relation to wing length, 4+5, and labrum;
number of ascoids on 3-15, or 16, and shape and relative length of that on 4; presence or absence of
papilla on 5); mandible (sharp or blunt tip, width and evenness of teeth); maxilla (shape, number of
lateral and ventral teeth, and dental depth, in sample specimen; palpal segment ratio; distribution of
Newstead’s spatulate sensilla, i.e. in most Old World species scattered or bunched on segment 3). Thorax:
scutal and pleural pigment; pleural hairs if present; inter-precoxal lobes (Sinton, 1928 : 745) if not
narrow as normal; wing (length and relation to width, R»/R2+3, R; overlap/R2); leg ratios, and femoral
spines if present. Abdomen: erect-hair sockets if present on tergites 2-6, best seen from above; sperma-
220
theca (shape and markings, collar if distinct, knob and surrounding pit, ducts and common duct if
present; furca and cercus if unusual).
$. Head: labrum length and its relation to wing length; cibarial teeth and pigment patch; pharynx;
antenna (as for 2); palpal ratio. Thorax: wing (as for 2); femoral spines if present. Abdomen: standard
taxonomic features of aedeagus, filaments and pump, paramere, coxite and its brush, and style.
FEATURES SOMETIMES USED. @: eye length, distance between eyes, interocular suture, arrangement of hair
sockets on head, markings on pigment patch, shape of cornua, hair sockets on basal costal node of wing,
nature of abdominal sternite 2. ¢: long antenna 1 and 2, large abdominal tergite 6, sternite 2, very long
hairs on terminalia, shape of surstyle. Early stages: seldom found in nature; various features can be
studied in eggs, larvae and pupae obtained by breeding. Patterns have sometimes been discernible on
eggs in gravid females, like that of S. jamesi.
Taxonomic features
The anatomy of sandflies has been described wholly or partly by Forattini (1973), Lewis (19735;
1975a), Perfil’ev (1968), Theodor (1958; 1965) and many others, and here it will suffice to mention
particular features.
The length of the labrum is taken as the distance from the tip of the clypeus (omitting mem-
branous connections) to the tip of the distal sensilla in the female, and to the tip of the incon-
spicuous small hood which covers them in the male. The labral sensilla of females are usually in
four groups, namely apical, subapical, adoral (along the shaft) and cibarial (mainly proximal to
the tip of the clypeus). Numerous variations include merging of adorals into subapicals or
cibarials, and differentiation of adorals into a fore and a submedian hind pair of rows. The
pattern of sensilla varies between many individuals and species but cannot practically be included
in all descriptions. The labral sensilla appear more or less equivalent to those of mosquitoes
(Lewis, 1975a), certain of which have some subgeneric significance.
The dorsal wall of the cibarium in the American Brumptomyia Franca & Parrot and Lutzomyia
Franga has a hind bulge which, according to Theodor (1965 : 173, 174), is unknown in Old
World species. Many of the latter, however, do have a bulge (Figs 117, 118, 181, 182, 186, 228-
231, for example) which, in S. knudseni at least, is less pronounced than in Lu. panamensis
(Shannon) (Fig. 232) and in the two species shown in sagittal section by Theodor.
The ventral wall of the cibarium between the chitinous arch and the arc of the hind teeth (here
called the inter-arcal area) varies in shape and is sometimes pigmented. The terms concave and
convex, applied to the cibarial and pharyngeal armatures, refer to their posterior borders.
The cibarium of most species of Phlebotomus either is unarmed, with merely scattered minute
spicules, or has scattered large and small spicules. Large ones are occasionally bigger than the
teeth of some Sergentomyia, which, however, are distinguishable by their linear arrangement. In
a few species, such as S. hassani and S. reidi, some teeth extend onto the cibario-pharyngeal
membrane.
The hind teeth in Sergentomyia, seen in ventral view, are often more or less foreshortened, and
in S. reidi, and possibly some others with pear-shaped teeth, the appearance is completely altered
by crushing.
In S. perturbans, S. reidi and some other species with a few strong teeth, the cibarial wall at
their bases is thick and refractive and probably acts like the base of a harrow supporting a row
of tines. The appearance is complicated in S. reidi by the presence of lines on the pigment patch
which, in sandflies, is nearly in the same focal plane as the teeth (Theodor, 1965 : fig. 1, AB).
In some females of S. barraudi the hind margin of the ventral cibarial wall has a row of lumps
which may be fore teeth displaced backwards.
The antennal ascoid formula is often used in males of Phlebotomus but varies in forms such as
P. longiductus, and one ascoid of a pair may be vestigial. The length of an ascoid on antenna 4
is sometimes a useful specific character (Parrot, 1940; Lewis, 1975), but (Schmidt & Schmidt,
1962) P. papatasi and P. argentipes (discussed later) show great variation. Antenna 5 bears a
papilla in most Phlebotomus but not in P. betisi or P. argentipes.
221
Occasionally hairs diverge widely from the shaft of the antenna (as in S. musai) or enwrap it
closely and obscure the joints so that the antenna looks like a thread (as in S. hamidi and S.
traubi).
The basal width of the main teeth of the mandibular armature (excluding extremities) may be
measured by dividing a length of about 15 um by the number of teeth in it. Teeth can be described
as small (about 0-9 um), normal (about 1-5 um as in S. antennata (Newstead)), or large (about
2:5 um as in S. fallax (Parrot)).
The base of the first maxillary palpal segment was taken as its junction with the blade. It and
other segments are often rather soft, and their apparent lengths depend somewhat on mounting
methods, but approximate measurements of some segments can be useful. Segments 2 and 3 have
distal knobs.
Wing length is a useful indication of the size of a fly. The length in female sandflies ranges
from 1-37 mm (a small S. barraudi from West Malaysia) to 3-8 mm (P. gigas Parrot & Schwetz),
and species can be grouped around the figures 1-5 mm, small, 2:2 mm, medium size, 2°7 mm,
large, and 3-6 mm, very large. The wings of males are shorter, and in one male of S. tiberiadis
from Saudi Arabia are only 1-17 mm long.
Lengths of the femur, tibia and basitarsus of each leg can be useful (Quate, 19626 : 252;
Quate & Fairchild, 1961 : 204; Quate & Rosario, 1962 : 788), and are here expressed as the
length of the femur followed by a ratio.
The femoral spines present in a few species come off easily and may be difficult to detect or
count in some specimens, because their sockets are rather like those of large hairs. If the legs
become detached before spines are examined the legs can be recognized as first, second or third
by the lengths of their tibiae.
The appearance of the spermatheca can vary according to the mounting medium used. The
internal ridges in S. dhandai are conspicuous in water but soon disappear in Berlese’s or probably
any other medium. The spermatheca of S. barraudi looks different in Fig. 64 and in Quate’s
(1962b) fig. 4d, and that of S. zey/anica in Fig. 204 and in Theodor’s (1938) fig. Vf and in Quate’s
(1962c) fig. 1f. In the latter instance a spermatheca of one type simulates another, and in the best
conditions it is sometimes difficult to differentiate a swollen tube from a narrow capsule. The
true shape may be obscured by disorientation, contraction or pressure from developing eggs,
and in many mounts the spermatheca is invisible. The collar is often a useful feature but may be
too indefinite. The spermatheca, an essential taxonomic character, must sometimes be carefully
studied in several individuals, preferably parous flies with no fat-body or developing eggs, or
gravid ones after detachment of terminal segments from the eggs.
The shape of the tip of the aedeagus is important but is sometimes intermediate between
blunt and sharp.
Classification
Classification of the Phlebotominae began in 1911 and some aspects are still debatable. The
system of Theodor (1948; 1958) is broadly. followed here.
The difference between Old and New World sandflies is real but difficult to define precisely
(Lewis, 1975a; Lewis & Lane, 1976). The significance of the dorsal wall of the cibarium, as shown
above, seems to be less clear than was thought. A very general difference between Old and New
World groups lies in the pattern of erect hairs on abdominal tergites 2-6. In the New World this
varies greatly (Lewis, 19755 : 366) but in the Old World there appear to be only two main
patterns.
Theodor’s subgenera of Sergentomyia have not been universally adopted, and even he evidently
regarded ‘Rondanomyia’ (Neophlebotomus) as not very well defined (1958 : 48), treated it with
some reservations, and included two exceptional species when dealing with the Palaearctic Region
only. The subgenera are used here because they avoid an unduly large assemblage of ungrouped
species, and facilitate faunal comparison with other regions. Some Oriental species of Sergento-
myia are easily placed in the subgenera Parrotomyia and Neophlebotomus, and others less easily.
222
Many, however, outside the distinct Sergentomyia, Grassomyia and Sintonius, form a miscel-
laneous category of species, some of which may be loosely connected with subgenera. Sub-
divisions of three subgenera are discussed later.
Considerable infraspecific variation is seen in the ascoids of P. argentipes, the cibarium of
S. barraudi, the maxilla of S. indica and various features of some other species. Divergent inter-
pretations have led to the existence of many synonyms, and to complex taxonomic citations of
species like S. ivengari. These become permanent features of catalogues, but may be reduced by
the use of informal names for little known taxa.
Key to the species and subspecies of Oriental Phlebotominae
The following key, like all keys, is based on a few of the many characters of each species, there-
fore for many species it is intended as an aid, and not a means, for identification. For some
sandflies, particularly some members of the subgenera Parrotomyia and Neophlebotomus, it is
necessary to examine several females and a male; the females, which are of more biological
interest, often have better characters, and a single one may have spermathecae shrunk or obscured
by developing eggs. These subgenera are difficult to define, and it is advisable to follow each in
the key in the case of problematical species.
The spermathecae provide important taxonomic features but those of genus Sergentomyia are
little used in the key because they are often difficult to see.
The key will require modification when some of the species are better known.
The following forms are omitted from the key. The female is unknown in P. nuri and newsteadi,
and S. A sp., B sp., Besout sp., brevinervis, displicata, kachekensis, morini, pooi, Rabok sp. and
Sepilok sp. The male is unknown in P. betisi, sejunctus, teshi and tubifer, and S. chakravarti,
cheongi, dayapensis (identity doubtful), dentacea, exastis, hamidi, jamesi, kauli, lagunensis,
mahadevani, modii, musai, Okinawa sp., pachystoma, spinifaucis, tonkinensis, tracheola and
yoshimotoi. Forms insufficiently described are S. angustipennis Q and 3), bigtii (3), nankingensis
(3), and torrechantei ($). These lacunae should not affect identification unduly because some of
the species are rare or local, and most gaps represent males of genus Sergentomyia in which most
determinations are based on the female.
1 Cibarial teeth absent or, if present, usually in the form of spicules and not in a definite row
(Fig. 27). Pigment patch nearly always absent. Hind ends of abdominal tergites 2-6 with
many erect hairs (Fig. 1), sockets as large as on 1. Style of male with three to five spines
(Figs 19, 25) and sometimes (some species of subgenus Jdiophlebotomus) with two hairs
as well (genus PHLEBOTOMUS, p. 233) ; 2
— Cibarial teeth in a transverse row (Fig. 65), fore teeth sometimes present (Figs 136, 245) and
usually pointing upward. Pigment patch (Figs 66, 136) usually present. Hind ends of
abdominal tergites 2-6 usually with all or most hairs recumbent, most sockets much
smaller than on 1 except in S. musai. Style of ¢ with four major spines and an accessory
seta (Fig. 80) (genus SERGENTOMYIA, p. 252). , : : 43
2 Cibarium with teeth. Pharynx unarmed. Palp not extending further than antenna 3;
Style of 3 with three to five spines, and a pair of intra-abdominal rods (subgenus
IDIOPHLEBOTOMUS, p. 250) ; , 3
— Cibarium with spicules or unarmed. Pharynx usually armed. Palp ex.ending further than
antenna 3 P : : : : : . : : : F ; : : 15
3 Female . : 2 ‘ ; ; f ; : < ° ‘ : : P 4
- Male ; ; i ' ; , ; : . i ; 11
4 Cibarial armature with median rod , 7 ; ; : ; : ; ; ‘ )
— Cibarial armature without median rod . ; ; ; : ; P 3 : : 9
5 Cibarial median rod with large serrations : ; : : : P. asperulus (p. 250)
— Cibarial median rod with minute serrations or none é s : : : : P 6
6 Cibarium without teeth except a few granulose spicules. ‘ ‘ . FP. erebicolus (p. 251)
- Cibarium withteeth . : : : , ; : : ‘ : : P , 7
7 Cibarial teeth all short - ’ ; : : : ; : , P. pholetor (p. 251)
— Some or all cibarial teeth long ; : , ; 8
223
24
25
26
Cibarial teeth in radiating lines : : ; ; ; P. tubifer (p. 252)
Cibarial teeth not in lines, very long, and parallel ; : : ’ P. frondifer (p. 251)
Cibarial teeth all small 3 F : : ; j 2 ‘ é P. stellae (p. 251)
Cibarial teeth not all small : : ; . ; ; : ; ‘ ; ; 10
Median tooth the longest . ; ; ‘ : ; : , : . P. teshi (p. 252)
Anterior teeth the longest . j ; 3 ; : P. sejunctus (p. 251)
Apical spine of style with marked basal expansion ; 2 ; . P. asperulus (p. 250)
Apical spine of style without marked basal expansion . : ; : : ‘ iZ
Style with three spines , ; : : n F ; ; P. erebicolus (p. 251)
Style with more than three spines s F : : ‘ ; : , 13
Style with four spines : , : : : : : ‘ ; P. frondifer (p. 251)
Style with five spines : : 14
Aedeagus prominent and capitate: paramere slender and without dorsal appendage
P. pholetor (p. 251)
Aedeagus small and triangular; paramere with basal dorsal curved appendage P. stellae(p. 251)
Coxite of 3 with sub-basal hairy process. Genital filaments 1-3-2:3 timesaslongaspump . 16
Coxite of ¢ without such process. Genital filaments 3-11 times aslongaspump . ‘ 24
Coxite 0:37-0:74 mm long, its process very small. Style long and cylindrical with three
terminal spatulate spines and two other spines. Paramere with two dorsal processes.
Surstyle with terminal spines. Pharyngeal armature comprising either a network of lines or
scales. Spermatheca with nearly equal segments and a refractive membrane (Figs 2, 3) near
bases of ductules (subgenus PHLEBOTOMUS, p. 233) : , 17
Coxite 0:20-0:33 mm long, its process usually large and having a brush et fone pairs: Style
not long, with four or five spines. Paramere simple, distal upper surface flat and elliptical
with short hairs. Surstyle without terminal spines. Pharynx of 2 with teeth or scales.
Spermatheca sometimes with differentiated rounded end-segment . ' : ; 19
Female. ; ‘ : , 2 P ; . P. papatasi (p. 233) ¢ or P. salehi (p. 235)
Male : 18
Upper Peers iobe fauch toners than paramere. eireeie with ee: or r Mometimes) hee
similar spines : : ; P. papatasi (p. 233)
Upper parameral lobe same enn as Saracens Surstyle aes seven spines, large to very
small . ; : . P. salehi (p. 235)
Style with five long spines, Ns at the ap and is near the middle Pines of 2 with irregular
scales or punctiform teeth (subgenus SYNPHLEBOTOMUS, p. 236) P. eleanorae (p. 237)
Style with four long spines, two near the tip and two near the base. Pharynx of 2 with large
backwardly directed teeth (Subgenus PARAPHLEBOTOMUS, p. 235) : i ‘ 20
Female . y ; d : ; ; : F : ; : ; : : 21
Male ; : : : : 22
Antenna 3 short (0- 12-0: 16 mm), 0- 5-0: 6 length of labrun ; : : P. alexandri (p. 235)
Antenna 3 long (0:23-0:33 mm), 0:7-1:0 length of labrum : ; P. sergenti (p. 236)
Basal process of coxite very large and thick with many hairs on its distal third P. nuri (p. 236)
Basal process of coxite small and thin with hairs only at its end Zo
Antenna 3 short (0:12-0:16 mm), 0-7-0:9 length of labrum. Genital pump 5 ction (0- 12 Sey
with small basal plate or funnel ; : P. alexandri (p. 235)
Antenna 3 long (0:25-0:34 mm), 1-0-1-4 length of ‘aera Genital pump long (0-17-0-2 mm)
with broad basal plate. , é P. sergenti (p. 236)
Style with four long spines of Bren one is enuinal, one Pabtee anal and two near the
middle.
Paramere with two or three lobes, with or without accessory spine. Aedeagus short and
conical. Pharynx with a small group of teeth in the middle and behind it some concentric
lines. Spermatheca segmented, yas segment not ee eee ANAPHLEBOTO-
MUS, p. 247) ; : : ; : 7 : 25
Style with five long spines . ; : : : . ‘ : : : ‘ ; 30
Female. ; c : : : ' , ‘ : i ; ; ; : 26
Male ; 28
Spermatheca slightly ‘carrot- shaped with small end-segment, individual duct about four
(possibly more) times length of spermatheca.
Sternal tubercle broad : ‘ b p : . : . P. colabaensis (p. 247)
224
38
39
Spermatheca spindle-shaped with very narrow cylindrical apical segment, duct short but
common duct very long.
Ascoids long. Palpal segment 3 with peg sensilla grouped around middle .
Pharyngeal armature with antero-median numerous long pointed teeth which blend laterally
with ridges. Individual ducts longer than spermathecae é : P. stantoni (p.
Pharyngeal armature with several antero-median rows of small short teeth, and antero-
laterally a number of backward-pointing teeth. Individual ducts shorter than sperma-
thecae . ; ‘ , , ; : ; F : : 2 P. hoepplii (p.
Paramere bilobed ; ; , ; 2 : : : ; P. colabaensis (p.
Paramere trilobed ,
Spine near aedeagus not longer than it. Pharynx with ¢ a series of oblique ridges radiating
from mid-line and ending in loops laterally i ‘ ; P. stantoni (p.
Spine near aedeagus much longer than it. Pharynx with a series of posterior ridges and,
antero-laterally, a number of teeth projecting medio-posteriorly . : P. hoepplii (p.
Paramere with three lobes (two in an Iranian species). Pharynx of 2 as in Anaphlebotomus
Spermatheca with differentiated end-segment (subgenus EUPHLEBOTOMUS, p. 240) .
Paramere without ventral process. Pharyngeal armature otherwise . d ,
Female
Male
Spermatheca with faint transverse striations : : : , P. kiangsuensis (p.
Spermatheca distinctly segmented
Spermathecal common duct with rather thin walls. ‘Antenna 5 without papilla P. mrcentines (p.
Spermathecal common duct with thick walls. Antenna 5 with papilla
Antenna 3/labrum 1-0 : f : j : : P. philippinensis phillapinensis (ee
Antenna 3/labrum 1-4 ; : ; P. philippinensis gouldi (p.
Middle lobe of paramere thicker (ana main fapoen) lobe : : iP; gj cst Le
Middle lobe of paramere thinner than main lobe .
Main lobe of paramere much more than twice length of middle lobe, lower lobe narrow,
depth of paramere about 0-29 of its length (measured to junction with coxite) P. argentipes (p.
Main lobe of paramere about twice or more length of middle lobe, lower lobe appearing
narrow but extending mesally, depth of paramere about 0°35 of its length
Antenna 3/labrum 1-7. Style 0-54 length of coxite, and 4-1 times as thick as Tone
P. philippinensis philippinensis (p.
Antenna 3/labrum 2:0. Style 0-61 length of coxite, and 3-6 times as thick as long
P. philippinensis gouldi (p.
Paramere truncated (not in subgenus) : : : ; ; . P. newsteadi (p.
Paramere not truncated
Pharynx of 2 and ¢ with punctiform teeth. ‘Spermatheca segmented, with long finger-like
process. Genital filaments 3—5 times as long as pump (subgenus LARROUSSIUS, p. 237) .
Pharynx of 2 with triangular or rounded group of medium-size teeth. Spermatheca incom-
pletely segmented. Genital filaments usually very oe 6:6-11-0 times length of pump
(subgenus ADLERIUS, p. sacs! ; ‘ ; . F ;
Female ‘
Male .
Pharynx with scarcely visible spicules '
Spermatheca with about 22 bead-like segments, a long neck and a very ‘small head.
West Malaysia ; ; : : : F : é . P. betisi (p.
Pharynx with readily visible spicules
Spermatheca with 30-35 segments, very long.
Median pharyngeal teeth larger than laterals : ; P. kandelakii burneyi (p.
Spermatheca with 8-21 segments.
Spermatheca nearly cylindrical, with 12-16 segments d ‘ ; P. major major (p.
Spermatheca narrowing at one or both ends, with about 18-21 segments P. keshishiani (p.
Aedeagus with mid-ventral surface finely serrated.
Aedeagus tapering gradually to a point through which genital filaments emerge
P. kandelakii burneyi (p.
Aedeagus smooth
Genital filaments 3—5 times length of pump : : : : . P. major major (p.
Genital filaments 6-11 times length of pump : : ; 5 . P. keshishiani (p.
225
46
65
66
Female. ; : ; P 4 P. chinensis chinensis (p. 239), P. longiductus (p. 240)
Male ; 47
Subterminal barb of aedeagus 30-35 vm from tip. Ascoid on antenna 4 about 0: 19 length of
segment ; ; P. chinensis chinensis (p. 239)
Subterminal barb of peda 12-14 um on tip. eco on antenna 4 about 0-31 length of
segment . . P. longiductus (p. 240)
Abdominal tergites 2— ‘6 with many erect hairs (musai-group, D. 253) : S. musai (p. 253)
Abdominal tergites 2-6 with few or no erect hairs : 49
Abdominal tergites 2-6 with a few posterior erect hairs (very ate in 3 S. Ciyded). Spermatheca
distinctly segmented (subgenus STNTONIUS, p. 307) : : 50
Abdominal tergites 2-6 usually with no erect hairs. Spermatheca not segmented boas some-
times striated or wrinkled F : : : : ‘ : , ; ; ; 63
Female . ‘ , : ‘ A , 5 A , , A . 3 ; 51
Male : 57
Pharynx very heavily armed. Cibarium with’ about 20 teeth i in n pallisade- like convex curve.
Two rows of fore teeth present : : ' : : , ; . S. orissa (p. 309)
Pharynx lightly armed or unarmed : : ; ; : é ‘ , ‘ 5 52
Pharynx with distinct spicules. : : : : : : ; S. hospitii (p. 309)
Pharynx with minute spicules or none : ‘ : ; A , 53
Cibarium with about 35 large pointed teeth in a row convex ipenaies ‘ S. eadithae (p. 309)
Cibarium with about 18 teeth or less. , ; : y ‘ , ‘ : ; 54
Cibarium with about five widely spaced teeth : : : ‘ S. christophersi (p. 308)
Cibarium with about 10-18 teeth close together . : ; : : : ’ ' a0
Cibarium without fore teeth : : 4 ; , : ‘ : S. sirohi (p. 310)
Cibarium with fore teeth . ' : , : F ; : ‘ 56
Cibarium with 12-15 long equal pointed teeth.
Pharynx with thick walls and an abrupt constriction behind the bulge. One to four papillae
on antenna 3, one or two on 4 F S. clydei (p. 308)
Cibarium with a comb-like row of rene 16- 18 seein fier ports directed upward and usually
hidden, outer teeth sloping towards centre. Pharynx thin-walled, narrowing gradually
behind bulge. One papilla on antenna 3 and 4 : : S. tiberiadis pakistanica (p. 311)
Cibarial teeth minute and arranged in groups : P . : , . S&S. clydei (p. 308)
Cibarial teeth otherwise. ; : : : : 2 : , ; ; . 58
Cibarium with about 28 teeth . ; : : : : ‘ ; S. hospitii (p. 309)
Cibarium with about 20 teeth or less. : , ; ; : : : 3 ? 59
Cibarium with two or three teeth : , : - : : . S. christophersi (p. 308)
Cibarium with ten or more teeth ; : : : ; : ; : 4 : 60
Cibarium with 20 teeth ; : : F g : , ; F S. eadithae (p. 309)
Cibarium with 14 teeth or less. ‘ 4 : : : : : ; : : 61
Antenna 3 = 0-06 length of wing ; : : : , : : . §. sirohi (p. 310)
Antenna 3 = 0-11 or 0-12 length of wing. ; F ; : : : ; : 62
Cibarial teeth on concave arc. A ' é ; F S. tiberiadis pakistanica (p. 311)
Cibarial teeth on arc convex in centre . c . ; : : ; . S. orissa (p. 309)
Spermatheca a round finely spiculate capsule.
Cibarium of 2 with convex comb of pointed teeth. Pharynx of 2 bulging near hind end.
Antenna of 2 with one ascoid on segments 4-15. Abdominal tergites 2-6 with some pos-
terior erect hairs. Genital filaments with rounded expanded tips. Parameres blunt (sub-
genus GRASSOMYIA, p. 268) : : : F : ; S. indica (p. 268)
Spermatheca elongate and usually without erieaies ‘ 64
Antenna 3 short, 0:08-0:19 mm, shorter than 4+5, usually shorter than Jabrum, Wing
narrow and lanceolate; Ro/ Ross usually less than one (0-3-0-8). Spermatheca tubular with
smooth sides and wide duct. Aedeagus thick. Style with four terminal spines, or two
terminal and two subterminal (subgenus SERGENTOMYIA, p. 253) . ‘ ; ; 65
Without this combination. Spermatheca usually not tubular ‘ : ‘ ‘ ‘ 69
Female. : 2 ; ' ; : : ; ; : , ; ; ¥ 66
Male 7 68
Hind teeth of pharyngeal armature much smaller rs fore teeth. Hind width of pharynx
about 0:58-0:77 of length ; , : P : ; f . JS. punjabensis (p. 255)
226
Hind teeth of pharyngeal armature not much smaller than fore teeth. Hind width of pharynx
about 0:37-0:59 of length
Length of pharynx 2-25 or less times hind width : - f : S. dentata (p.
Length of pharynx 2:26 or more times hind width. ; , S. theodori pashtunica (p.
Parameres hooked . ‘ : : ; . iS. punjabensis (p.
Parameres blunt : ‘ : S. Wentisa (p. 253), 'S. theodori pashtunica (p.
Cibarial armature usually a Comb: like row of parallel teeth with short points. Pharynx often
lamp-glass-shaped. R./R23; often 0-3-1-:0 (in some Palaearctic species). Spermatheca a
smooth, round or elliptical, capsule. Aedeagus slender, triangular and narrowing gradually,
usually to a sharp point. Paramere hooked. Style with all or two of four spines terminal
(subgenus PARROTOMYIA, p. a ;
Without this combination .
Female
Male
Pharynx with many distinct pointed teeth
Pharynx with very fine spicules or none
Cibarium with notch in hind end of ventral plate, and 10-50 teeth i in concave row
Cibarium without notch in hind end of ventral plate
Cibarium with about 10-14 teeth, notch shallow . : ; ; Ss. shorttii (p.
Cibarium with about 24-50 teeth, notch deep 3 ; . : . : :
Cibarium with 24-34 teeth : ; j : , : . IS. babu babu (p.
Cibarium with 45-50 teeth. (Sri Lanka) : : p : : S. babu insularis (p.
Spermatheca nearly spherical.
Cibarium with 20-22 teeth in a straight line. care : ; . S. palestinensis (p.
Spermatheca not nearly spherical :
Pharynx broad, often with rather straight sides, vith many iene Apely outed teeth, Gibariut
with 40-70 teeth and tip of pigment ie bifid, ae orfenestrated . S. barraudi (p.
Without this combination
Cibarium with 50-60 short teeth in row concave in ene ad Siaight 6 or convex at Bee
fore teeth in two rows. , : 5 : , ; : . IS. spinifaucis (p.
Without this combination .
Cibarium with 64 teeth or more
Cibarium with 60 teeth or less
Pharyngeal teeth long, numerous and finely pointed. Cibarial teeth 1 more than 65
Pharyngeal teeth not long .
Palpal segment 4 = 1-6 times length of a Cibarium with 65- 10 hind teeth
Antenna 3 = 0:28-0:31 mm long. Cibarial fore teeth in two rows of about 22. (Philip-
pines) . , S. mangana a
Palpal segment ee = Ls 1 times length of ef Cibarium with 90 hind teeth
Antenna 3 = 0:11-0:14 mm long. Cibarial fore teeth numbering 35, in two ill- denied rows.
(Australia and Nusa Tenggara) ’ S. queenslandi meridionalis (p.
Antenna 3 = 0:26-0:33 mm a Cibarial ae jest in fae rows of about 22. (West Malay-
sia) : ; ‘ i p : , S. rudnicki (p.
Hind margin of pigment patch concave ; : : . S. himalayensis (p.
Hind margin of pigment patch convex. (South Vietnam) 2 : S. brevicaulis (p.
Cibarium with 10-12 teeth. (Philippines) : : : ‘ ; . . S. bigtii (p.
Cibarium with 26 or more teeth ; ‘ , ‘ ; : , d ;
Cibarium with 42-50 teeth ‘ ; : : A : ‘ S. africana magna (p.
Cibarium with 26-32 teeth ; : ; : é : , . : ‘ :
Cibarial teeth on a distinct arc. (India) . : F : ‘ S. kauli (p.
Cibarial teeth on a nearly straight line. (Philippines) : i : S. torrechantei (p.
Cibarium with deep notch in hind end of ventral plate.
Cibarium with 16-20 teeth. Pharynx with transverse ridges and some hind spicules
S. baghdadis (p.
Cibarium without such notch . : -
Pharynx with well-defined scales. (Pakistan) F ° mS grekovi (p.
Pharynx without scales but sometimes with a few transverse wrinkles or minute spicules
Cibarium with 60-90 teeth
Cibarium with 34 teeth or less
227
111
112
Cibarium with about 90 teeth, very narrow and almost invisible. Pharynx lamp-glass-shaped,
without spicules. (Borneo) : ; . §&. denticulata (p.
Cibarium with about 60 teeth, easily visible. Pharynx with nearly straight sides and a group
of hind spicules. (Nusa Tenggara) . ; ; ‘ ; ‘ ; S. timorica (p.
Cibarium with 24-32 hind teeth
Cibarium with 14-18 hind teeth ; ; : : ; ; ‘ ‘
Cibarial teeth on convex arc : ; ; : é : : S. franciscana (p.
Cibarial teeth in nearly straight line ; : 2 : F : : ;
Cibarium with about 24 teeth . : ; , ; : : . &. dayapensis (p.
Cibarium with 30-34 teeth : f P : . S&S. bukidnonis (p.
Cibarium with 17 hind teeth and a pigment patch f . F : . S. modii (p.
Cibarium with 14-18 hind teeth and no pigment patch
Hind teeth in convex row. Spermatheca ovoid with protuberant knob. (Philippines)
Tip of aedeagus pointed
Tip of aedeagus rounded
Style with four apical spines
Style with two of spines subapical
Cibarium with an irregular row of 13 fore teeth S. ‘brevinervis (p.
Cibarium with no fore teeth or with a straight row of a few at bases of hind teeth
Cibarium with about 20-30 hind teeth ; F : : ; S. africana magna (p.
Cibarium with about 60 hind teeth . : : ; S. queenslandi meridionalis (p.
Cibarium with 16 or 17 teeth
Fore teeth absent, pigment patch faint. Pharynx with narrow hind end S. franciscana (p.
Cibarium with about 19 or 20 hind teeth
Cibarium with two rows of fore teeth and a long narrow 7 picinert etc S. brevicaulis (p.
Cibarium with no fore teeth and no visible pigment patch : S. denticulata (p.
Style with three large and one small spines ‘ ‘ : : , S. himalayensis (p.
Style with four equal spines
Style about five or six times as long as thick.
Style about four times as long as thick
Pharynx unarmed
Pharynx armed
Cibarium with 12-17 hind teeth and S1x x fore teeth. : , S. bukidnonis (p.
Cibarium with about 30 hind teeth, fore teeth faint or absent . : S. timorica (p.
Cibarial fore teeth well developed : ; ) : ; : ; S. grekovi (p.
Cibarial fore teeth faint
Antenna 3 = 0:29-0:38 mm long ; , : : : : S. rudnicki (p.
Antenna 3 = about 0:16-0:18 mm long
S. babu babu (p. 257), S. babu insularis (p. 258), S. baghdadis (p. 258), S. shorttii (p.
Two of spines on style subapical : ‘ , iS. sp. (A) (p.
All spines on style apical : : ; ; : ; : ‘
Cibarium with about 18 teeth . : ; p : , : ‘ Ss. atau (p.
Cibarium with about 10-13 teeth : : : : : ; 2 , :
Pharynx unarmed . : : : 3 : : : : ’ . S. sp. (B) (p.
Pharynx armed : ‘ : ‘ : e : ; ; ; :
Pharynx with pointed teeth : : , 3 , . : , S. mangana (p.
Pharynx with scale-like ridges. : , S. palestinensis (p.
Cibarial tooth-row usually comprising parallel iecth, ohien neatly equal but not very narrow.
Pharynx slender, with teeth or scales, or nearly unarmed. Antenna 3 long, longer than
4+5, often 1:25-2:00 times length of labrum. Wing rather broad, R, usually longer than
Rz43. Spermatheca often a thin-walled capsule, sometimes with cross-striations; duct
sometimes narrow and joining common duct. Aedeagus usually slender with blunt tip.
Paramere hooked. Style with two terminal spines and two others more or less terminal or
near the middle (Subgenus NEOPHLEBOTOMUS, p. ee
Without this combination . : ,
Female
Male :
Cibarium with about eight 1 rows of fore teeth
228
S. yoshimotoi (p.
111
155
112
134
113
Cibarium without eight rows of fore teeth
Cibarium with fore process of pigment patch absent or ‘faint. Labrum 0: 18-0- 21 length of
wing. : S. gombaki (p.
Cibarium with ‘fore process of Brement patch maornnant Cabra 0:13-0:15 length of
wing . : : : : ; : . S. arboris (p.
Cibarium with (hres rows of fore fecih’
Cibarium with less than three rows of fore teeth or none
Fore teeth of hind row very large ; , , : : ; : S. gemmea (p.
Fore teeth of hind row not very large . : ; : .
Labrum very long, 0-18—0-20 length of wing. Rik eS = re 53-2: 06 : S. malayae (p.
Labrum not very long, 0:13-0:15 length of wing. R2/R2,3 = 2:10-2:85. S. zeylanica (p.
Cibarial central teeth markedly different from the rest : 5 : : :
Cibarial central teeth not markedly different from the rest
Cibarial central teeth much larger than the rest
Cibarial central teeth not much larger than the rest
Antenna 3 = 1:34-1:42 length of labrum : : : ; : S. silvatica (p.
Antenna 3 = 2:19-2:36 length of labrum ‘ : : : , ;
Wing length 2:15-2:23 mm. Spermatheca with simple duct , ; : . S. quatei (p.
Wing length 1:79-1:90 mm. Spermatheca with modified duct . ; ; S. hamidi (p.
Cibarium with 24 teeth : : : : ; , ; : : S. dhandai (p.
Cibarium with 17 teeth or less
Cibarium with 14-17 hind teeth, fore teeth absent. or varying from | one row of four to two
rows of up to 20; pigment patch with anterior ee a either thick, small or absent
(leaving patch hemispherical) : : ; : S. iyengari (p.
Cibarium with about 11 teeth, and one enue row of fore ei Pigment patch with broad
anterior projection . : : ‘ , , : S. tambori (p.
Cibarial pigment patch very narrow and linear : ; : ; g S. linearis (p.
Cibarial pigment patch not very narrow
Cibarium with 18 fold-like teeth longest in the centre
Antenna 3/labrum 2-0. Spermatheca an oval sia with faint streaks . S. traubi (p.
Cibarial teeth not fold-like
Cibarium with 14 hind teeth, of which the lateral one on each side i is ‘separated from the rest,
and about ten fore teeth in one row; pigment patch with marked anterior projection and
posterior notch E ‘ : : : . S. nankingensis (p.
Cibarial teeth and pigment patch not like this , : : ‘ :
Cibarium with nine or ten groups of denticles ; : : : . iS. tonkinensis (p.
Cibarial teeth not in such groups
Cibarium with 50—60 teeth : : ; 2 . S. hodgsoni hodgsoni (p.
Cibarium with about 20 teeth or less
Cibarium with about 20 teeth :
Cibarium with less than 20 teeth on average
Cibarial teeth on a straight line . ; : 4 ; ; ; : . S. balica (p.
Cibarial teeth on a curve
Cibarial teeth contiguous, in row convex in centre , : : : . S. khawi (p.
Cibarial teeth separated, in concave row
Pigment patch narrow and long . : F . S. purii (p.
Cibarium with eight or nine main hind teeth fice from fetiactive brow area with colour
different from pigment patch : : ; : ; , . JS. perturbans (p.
Cibarium otherwise
Cibarial teeth separated.
Seven to 15 of them ina regularrow . : ‘ j , : S. jefferyi (p.
Cibarial teeth contiguous
Cibarium with about 14 hind teeth i ina ‘line angular at the centre; seven round teeth present
behind hind ones. . S. chakravarti (p.
Cibarium with about eight hind teeth i in nearly straight row; no teeth behind hind ones
S. malabarica (p.
Style with two of spines near middle
Style with spines terminal or subterminal
Brush on coxite with about 183 close hairs. Style not very swollen near t middle S. gombaki (p.
229
114
275)
270)
115
17
273)
116
282)
292)
118
123
119
121
290)
120
288)
275)
271)
122
77)
291)
280)
124
292)
125
282)
126
291)
L227
277)
128
129
131
270)
130
278)
288)
283)
132
278)
133
271)
280)
135
146
275)
Brush on coxite with about 95 hairs or less, or indefinite
Brush on coxite with about 95 close hairs. Style very wide near middle S. sp. (Besout) (p.
Brush on coxite with 60 hairs or less, or absent
Cibarial teeth fold-shaped . ; ; ; . : : : Os franbi (p.
Cibarial teeth not fold-shaped
Style with seta at about 0:8
Style with seta at 0-7 or more proximal
Coxite with long brush of about 60 hairs. : ‘ : ‘ ; . S. purii (p.
Coxite with a short brush of about 31 hairs . : : 5 : : S. quatei (p.
Coxite long and narrow and slightly curved . ; : ’ , ee dis pertarbans Me
Coxite not long and narrow and slightly curved :
Aedeagus thick, mid width of shaft about 0-19 of extreme length of aedeagus 7
Aedeagus slender, mid width of shaft about 0:11 of extreme length of aedeagus
R,/R2,3 about 1:2. Coxite with seta distinctly proximal to middle spines . S. khawi (p.
R,/Rz.3 about 1-5—2-7. Coxite with seta near middle spines 4 F :
Coxite with some of outer hairs concentrated ; : : ; 4 S. deylanica (.
Coxite with outer hairs evenly spaced é . ; ; S. malayae (p.
Cibarial fore teeth about four deep in broad ‘band 2 : 3 S. sp. (Rabok) (p.
Cibarial fore teeth not in broad band . ; 2 A ; ‘ , : :
Brush starting at 0-21 of coxite ; : : ; ; : A S. arboris (p.
Brush starting at 0-23 of coxite : : : : F S. gemmea (p.
Style with spines of unequal thickness, three ‘terminal.
Coxite without definite brush : : : : S. tambori (p.
Style with spines of equal thickness, two or four ante
All spines of style terminal ‘
Two spines of style at 0-76-0-85 .
Paramere with spinose process at base of neck : F : . S. hodgsoni hodgsoni (p.
Paramere without such process ‘ ; , , : : : : ;
Cibarium with long narrow pigment patch . ; ; : ; : S. linearis (p.
Cibarial pigment patch not long and narrow : : ; : ‘ , ‘
Cibarium with conspicuous fore teeth ; ; ; ; : ; S. iyengari (p.
Cibarium without fore teeth : 5 ; P S. dhandai (p.
Cibarial hind teeth comprising four central ites ones ad a row of about five small ones on
each side : : , : , : , : : , S. silvatica (p.
Cibarial teeth not like this .
Cibarium with about eight teeth, some ° with several points, and a ‘few vestigial fore teeth
S. balica (p.
Cibarium otherwise ; ; , : p : : :
Cibarium with about six hind and about Six fore teeth : : : S. sp. (Sepilok) (p.
Cibarium otherwise : : : : : A
Cibarium with ten peattened Hint teeth and i no fore teeth 4 : . iS. malabarica (p.
Cibarium with about seven ill-defined hind teeth, not widely scattered, and about 15 small
fore teeth : , ; , S. jefferyi (p.
Cibarium of 2 with one or more rows of small, sometimes scarcely visible, hind teeth.
Spermatheca smooth, an elliptical or cylindrical ‘capsule. Aedeagus pointed and paramere
hooked. Style with all spines terminal or two of them subterminal (nicnic-group, p. 294)
Without this combination (not grouped, p. 295)
Female
Male : ; ; ,
Hind end of pharynx with ¢ a group of small but conspicuous dark spicules . S. nicnic (p.
Hind end of pharynx without such spicules.
Cornua divergent : F ; 5 : , ; > . S. bailyi (p.
Cibarium with multiple row of teeth : : : : : : : . S. bailyi (p.
Cibarium with single row of main teeth : 5 ; . : ; : ‘
Cibarial hind teeth diamond-shaped . : 7 . : ; S. kachekensis (p.
Cibarial hind teeth not diamond-shaped ; ‘ : j : " .
Pigment patch present. Antenna 3 about as long as 4+ 5 ; : , . §&. nicnic (p.
Pigment patch absent. Antenna 3 longer than 4+ 5 A ; , . S. displicata (p.
Female : : : ; , i , : : :
230
Male
Cibarium without row ’ of teeth : ‘ , ; ; g : ; US: jamesi :
Cibarium with row of teeth
Cibarial teeth (in the usual ventral view) broad and often diamond- shaped o or pear-shaped
Cibarial teeth not broad except at bases
Cibarium with cluster of small teeth behind main teeth . i 5 Ss. reidi (p.
Cibarium without such cluster
Cibarium with eight wedge-shaped teeth.
Pigment patch absent. Pharynx rather slender with compact group of teeth S. tracheola (p.
Cibarium with pear-shaped teeth
Cibarium with about 25 teeth rather like fish hooks S. maai (p.
Cibarium with 18 teeth or less
Cibarium with 12-18 hind teeth ; : , , 7 : ; S. heiseri (p.
Cibarium with eight to ten hind teeth : : : ;
Cibarial arch about 4-9 tooth lengths from teeth, intervening area dark : S. losarcus (p.
Cibarial arch about 1-6 tooth lengths from teeth, intervening area not dark S. cheongi (p.
Cibarium with long median projection ; i ; : : ; S. anodontis .
Cibarium without such projection
Cibarium with 38 very long teeth in convex row and 65 fore teeth in seven or r eight rows
S. dentacea (p.
Cibarium without this pattern
Cibarium with 70 teeth. Pharynx with ten short spicules. "Ro|Ross = 4: 8. Spermatheca
subovoid ; ‘ ‘ , ; : ; : ; S. sp. (Okinawa) (p.
Without this combination :
Cibarium with several rows of lateral teeth.
Pigment patch with broad process and narrow hind part. Pharynx narrow with many
teeth . . : f A : 3 SS. montana (p.
Cibarium without several rows of lateral teeth :
Cibarium with 14 inwardly sloping teeth, and no fore teeth or Ppnent atte (philippines)
S. exastis Me
Without this combination .
Cibarium with 30 short linear teeth in compact row and about 80 fore teeth i in five or six rows.
Pharynx with dense patch of teeth. Antenna 3 about twice length of labrum. (Philippines)
S. dapsilidentes (p.
Without this combination . :
Cibarium with 10-14 teeth looking like barbed fish hooks: fore teeth absent: pigment patch
rectangular. Pharynx unarmed. (Philippines) . : : ‘ S. delfinadoae ae
Without this combination .
Cibarium with many small triangular ‘teeth of different sizes ; tending to form three TOWS
centrally. Pharynx with finely spiculate act en ‘ ; S. fanglianensis (p.
Without this combination .
Pharynx distinctly armed ;
Pharynx unarmed or with minute spicules
Cibarium with 26 teeth . : : : : ; P . oS. lagunensis (p.
Cibarium with 50 teeth : / , : : : ' ; . §&. mahadevani (p.
Antenna 3 = 0-12 mm long : : : Z : ‘ ; ’ . SS. neras (p.
Antenna 3 = 0:19-0:54 mm long
Cibarium with about eight small blunt teeth in a neaelG meen ie ; S. hassani (p.
Cibarium with 9-29 pointed teeth on a distinct arc
Cibarium with many fore teeth
Cibarium with very few or no fore teeth
Cibatium with 10-12 teeth pe ae et Os Na Saimin ip:
Cibarium with 20-29 teeth
Interarcal area sclerotized and wrinkled. ‘Antenna 3 about 1:3 length of labrum, ascoids
with spur. Tergite 8 with patch of hairsoneachside . : ‘ . iS. pachystoma (p.
Cibarium with nine teeth . : ; . d ; ; : : S. kelantani (p.
Cibarium with 12-27 teeth : : . 2 : : ‘ ; ‘ ;
Cibarium with 12-15 teeth : : ; ; ; : ‘ 3 S. hitchensi (p.
Cibarium with 24-27 teeth ' : : : ; : ; ; S. knudseni (p.
231
185 Cibarium with patch of teeth behind main teeth . ; ; ' : . IS. reidi (p. 305)
— Cibarium without such patch 186
186 Style with two of spines subterminal and seta 1 near base, and cibarial teeth small S. hitchensi (p. 299)
— Without this combination . : : : 187
187 Cibarium with faint median hind process and no definite teeth ; , Ss. anodontis (p. 295)
— Cibarium otherwise . ; i : : : ? ; ; . 168
188 Palpal segment 4 nearly twice length of 3
Cibarium with 12 short linear teeth and ten fore teeth in two rows _ S. dapsilidentes - 296)
— Palpal segment 4 not so long ? 189
189 Cibarium with six to ten barb-like teeth, six fore teeth and a long antenna 3, 0- 31 mm
S. acted (p. 296)
— Without this combination . 190
190 Cibarium with many small triangular teeth of different sizes tending to form two rows in the
centre ; ; : ; ; : : S. fanglianensis e 297)
— Cibarial teeth not hice Gis. : P 191
191 Cibarium with eight prominent diamond- shaped hind teeth and eight fore teeth i in two rows.
Inter-arcal area pigmented ; ; ; F ; : , . S. heiseri he 299)
— Cibarium not like this ; 192
192 Cibarium with about eight sharp spike- like teeth, and 10-20 fore en usually i in one main
row. , : : : ; ' ; : : ; . JS. imitor (p. 299)
— Cibarium otherwise : : 193
193 Cibarium with about eight distinct separated teeth o on convex arc, a strong bulge, and pigment
patch about two-thirds width of cibarium p . , ‘ : S. knudseni (p. 300)
— Cibarium otherwise . : 194
194 Cibarium with seven to nine pear-shaped teeth, no fore teeth and pigmented inter-arcal
AalCa oe : : P : : : : é , ‘ S. losarcus (p. 301)
— Cibarium otherwise ; : : 4 : ‘ : . 195
195 Cibarium with 12-15 small teeth and eight fore teeth ; : : : . S&S. maai (p. 301)
— Cibarium otherwise . : 196
196 Cibarium with about 18 teeth in one row, about three rows in front of them on each side, and
a distinct pigment patch . : : E , , . : ; S. montana (p. 303)
— Cibarium otherwise . 5 197
197 Cibarium with six or seven groups of small fine teeth, and some fore teeth, and no pigment
patch . ; ; , : : 2 ; : ; . ; . iS. morini (p. 303)
— Cibarium otherwise . 198
198 Cibarium with six small but distinct teeth, five fore teeth i in one row, and: no pigment patch.
Antenna 3 short, 0-11 mm, style with three apical and one subapical spines, and seta at 0-7
S. neras (p. 303)
— Without this combination . : : : . &99
199 Cibarium with about six long teeth and about 18 irregular fore teeth . . S. pachystoma (p. 304)
— Cibarial hind teeth numbering 23-29; fore teeth numbering 10-15, those at centre less
pointed than side ones. , : : ; : ' ; ; . IS. pooi (p. 305)
Taxonomy and distribution of species
Under each genus and subgenus references to descriptions are cited and brief notes are given on
some characters and on distribution. Diagnostic summaries are included in the key.
The species in each genus are arranged in alphabetical order. The taxonomic citations include
all references to Oriental species, except some out-of-date ones, and a few to species in other
regions. Where holotypes were not studied, paratypes were often examined. Descriptions are
given for new species and a number of others.
Specimens headed ‘Material examined’ were studied in detail. Many others were identified or
checked, and some of them are indicated by italicized names of collectors in the distribution lists.
Some localities are omitted, either because they cannot be traced, or because they are near other
recorded localities, or because they were recorded before 1928 when few species of Sergentomyia
were reliably identified. Table 2 gives the position of localities not shown on The Times Atlas of
1972. Map | shows the sources of all the material referred to in this study.
Zoe
Genus PHLEBOTOMUS Rondani & Berté
Flebotomus Rondani & Berté in Rondani, 1840 : 12. Type-species: Bibio papatasi Scopoli, by monotypy.
Phlebotomus Rondani & Berté [emendation]; spelling fixed under suspension of rules by ICZN, 1954,
Opinion 256 : 199; Theodor, 1948 : 96; 1958 : 16; 1965 : 179; Lewis, 1967 : 14.
Phliebotomus subgenus Phlebotomus Rondani & Berté; Quate, 1964 : 238.
Normally there is no row of cibarial teeth and no pigment patch. On the antenna there is a
papilla on segment 5 and, in the male, two ascoids on segments 3-15. Erect hairs are present on
abdominal tergites 2-6, and there are four or five spines on the style of the male. The genus is
widespread in the Old World, and most species occur in the north.
Subgenus PHLEBOTOMUS Rondani & Berté
Phlebotomus subgenus Phlebotomus Rondani & Berté in Rondani, 1840 : 12; Theodor, 1948 : 96; 1958 : 16;
Perfil’ev, 1968 : 227; Hennig, 1972 : 53.
The pharynx bears ridges or scales and the spermathecal segments are equal. The genital filaments
are short, each paramere bears two processes, the long coxite carries a lobe and the style is long.
The subgenus is represented in the west of the Region.
Phlebotomus (Phlebotomus) papatasi (Scopoli)
(Figs 1-5, Map 2)
Bibio papatasi Scopoli, 1786 : 55. No types mentioned: ITALY.
Phlebotomus papatasii; Howlett, 1915: 294 [misspelling]; Sinton, 1924a: 814; 1925a:468 [surstyle
variation etc.]; 1925d: 107; 1927d: 27; 1928c : 300 [synonymy]; 1932a:59; 1933d: 418; Sinton &
Barraud, 1928 : 329; Mukerji, 1931 : 442 [larva]; Mitra, 1952 :550 [palp sensilla]; Mitra & Mitra,
19536 : 434.
Phlebotomus papatasi (Scopoli); Schmidt & Schmidt, 1962 : 723.
Phlebotomus (Phlebotomus) papatasi (Scopoli); Parrot, 1940:310; 1946:67; Theodor, 1948 : 106;
1958: 17; Quate, 1964: 240; Lewis, 1967: 14; Perfil’ev, 1968 : 228; Abonnenc, 1972:99; Bhat &
Modi, 1976 : 265, 266.
Recent full or partial descriptions were given by Abonnenc, Perfil’ev, Quate and Theodor.
In the male there are two, and sometimes three, spines on the surstyle, the dorsal process of
the paramere carries hairs only ventrally and is much longer than the rest of the paramere, and
the first two spines of the style are close together. The pharyngeal teeth of the female form a
wide-meshed network.
The proximal part of the spermatheca (near the ductules) has been variously described because
its structure is difficult to see. Comparison of specimens in Berlese’s medium and in water with
Theodor’s (1965 : 174) figure shows the following features. The chitinous capsule comprises
about nine segments or hollow annuli, the most proximal of which is small and collar-like. The
head has a thick lateral wall and is connected to the long ductules, the bases of which are obscured
by a mushroom-like halo of refractive material which is part of the outer covering of the sperma-
theca.
2 (extra facts). Labrum with upper brush-hairs blade-like, two median apical sensilla long and narrow
and adorals small. Hypopharynx with about 16 teeth on each side. Maxilla with about six lateral and 17
' ventral teeth, and a dental depth of 0-10 mm, clavate sensilla concentrated near middle of segment 3 of
palp.
MATERIAL EXAMINED.
Pakistan: Taxla, 1 9.
DisTRIBUTION. Bangladesh: Parbatipur, Rajbari (Sinton’s notes). India: Pusa (Annandale, 1911c : 320;
Barraud, 1926 : 214; Craighead & Das, 1928 : 862; Howlett, 1915 : 296); Howrah (S. Das); Osmanabad
(V. Dhanda); Aurangabad, Jalna, Patan (Farooq & Qutubuddin, 1945 : 85); Baramula, Ramban, Riasi,
Srinagar (Jacob & Kalra, 1951 : 325); Tibi (N. L. Kalra); Jalor (Kaul et al., 1973 : 532); Jammu, Uri
239
SW
WY
Figs 1-9 Phlebotomus species. 1-5, P. papatasi, 2: (1) abdominal tergites 2-6; (2) spermatheca in
water; (3-5) same in Berlese’s medium. 6, P. betisi, 2, labrocibarium. 7, P. kandelakii burneyi, °,
labrocibarium. 8, P. keshishiani, 2, labrocibarium. 9, P. major major, 2, tip of labrum.
234
(Mitra, 1953a : 324); Hoora (Mitra, 1953b : 158); Khandwa, Mahad, Mahableshwar, Mundwa, Panch-
gani, Poladpur, Wai etc. (Mitra, 1954a: 111; 1955 : 82); Nowshera in Kashmir (Mitra, 1959 : 59, 62);
Hyderabad (commonest species, Qutubuddin, 1944 : 208); Kotelanka, Undi etc. (R. Reuben); Bikaner
(Sharma et al., 1973c); Kirki, Poona (Sinton, 1924f : 1042); Bombay, Maini Majera, Mandapam, Melur,
Nagpur, Pamban Island, Roorkee, Saharanpur etc. (Sinton, 1927b : 942); Calcutta, Madras (Sinton,
1932a: 70); Sagar (Sinton, 1932c : 577); Agra, Ajmer, Anandpur area, Coimbatore area, Dehra Dun,
Ferozepore, Lucknow, Madhopur, Mahendragarh, Mescara, Naini Tal area (Sinton’s notes). Pakistan:
Abbottabad, Bannu, Chaman, Chilas, Dera Ismail Khan, Digri, Gilgit, Hyderabad, Idak, Jamesabad,
Jamrud, Jandola, Jhelum, Kandhkot, Karachi, Kashmore, Khairpur, Khirgi, Kohat, Lahore, Landi
Kotal, Larkana, Miramshah, Mir Muhammad, Mirpur Kas, Nowshera, Pano Aqil, Peshawar, Quetta,
Rawalpindi, Saidpur, Shikarpur, Tando Bago, Tank, Taxla (Lewis, 1967 : 14); Kotkai, Multan (Sinton’s
notes).
According to Sinton (1932a : 70) this species was scattered all over the plains of India, especially
in hot dry areas, and was found as far east as Calcutta and as far south as Madras, but was
commonest in the north-west. He (1925d : 703) did not find it above 610 m in eastern India,
probably because the monsoon makes the hills too wet.
Phlebotomus (Phlebotomus) salehi Mesghali
(Map 2)
Phlebotomus (Phlebotomus) salehi Mesghali, 1965 : 264; Mesghali & Rashti, 1968: 770 [9]; Kalra &
Lewis, 1976 : 522. Holotype 3, IRAN (depository not stated) [not examined].
In the male the dorsal process of the paramere is shorter than in P. papatasi, and the surstyle
bears seven spines ranging from large to very small.
2 (extra facts). Labrum and maxillary sensilla as in P. papatasi. Hypopharynx with 17 teeth on each side.
Maxilla with six lateral and 17 ventral teeth and a dental depth of 0-08 mm.
MATERIAL EXAMINED.
India: Tibi area, 1 9.
DisTRIBUTION. India (Rajasthan): Bikaner and Tibi areas (Sharma et al., 1973a; 19736; Kalra & Lewis,
1976).
Subgenus PARAPHLEBOTOMUS Theodor
Phlebotomus subgenus Paraphlebotomus Theodor, 1948 : 97; 1958 : 19; Perfil’ev, 1968 : 232. Type-species:
Phlebotomus sergenti Parrot, 1917, by original designation.
The sub-basal process of the coxite is medium-sized to large and bears long hairs, and the style
has four spines. This subgenus occurs mainly in the Palaearctic Region where it includes some
closely related species. It is represented in the west of the Orient.
Phlebotomus (Paraphlebotomus) alexandri Sinton
(Map 3)
Phlebotomus sergenti var. alexandri Sinton, 1928c : 308. Type not indicated, PAKISTAN (depository
unknown) [not examined ].
Phlebotomus alexandri Sinton; Sinton, 1932a : 58; 1933e : 418.
Phlebotomus (Phlebotomus) alexandri Sinton; Parrot, 1940 : 310; 1946 : 67.
Phlebotomus (Paraphlebotomus) alexandri Sinton; Theodor, 1958 : 19; Theodor & Mesghali, 1964 : 290;
Lewis, 1967 : 15; Perfil’ev, 1968 : 241 [variation].
The authorship and date of P. alexandri are Sinton, 1932: 58 (ICZN, 1964: Article 10(b)).
Perfil’ev and Theodor have given recent descriptions.
In the male the basal lobe of the coxite is thickened at the end and antenna 3 is short. The
pharynx of the female is conical. This small sandfly with rather narrow wings looks rather like a
species of Sergentomyia.
235
2 (extra facts). Labrum and palp sensilla as in P. papatasi. Hypopharynx with about 16 teeth on each side.
Maxilla with four lateral and nine ventral teeth, and dental depth of 0:06 mm.
MATERIAL EXAMINED.
Pakistan: Qambar.
DISTRIBUTION. Pakistan: Dehra Ismail Khan, Hyderabad, Kandhkot, Larkana, Parkuta, Qambar,
Shikarpur, Tank (Lewis, 1967 : 15).
Phlebotomus (Paraphlebotomus) nuri Lewis
(Map 3)
Phlebotomus (Paraphlebotomus) nuri Lewis, 1967 : 15; Artemiev, 1974a : 160, 161. Holotype 3, PAKISTAN
(BMNH) [examined].
P. nuri is the only Oriental species with a large lobe on the coxite.
DISTRIBUTION. Pakistan: Rawalpindi, Said Pur (Lewis, 1967).
Phlebotomus (Paraphlebotomus) sergenti Parrot
(Map 3)
Phlebotomus sergenti Parrot, 1917: 564; Sinton, 1924a:814; 1928c:307 [synonymy]; 1932a: 58;
1933e : 418; Sinton & Barraud, 1928 : 329. Syntypes 3, ALGERIA (depository unknown) [not examined].
Phlebotomus (Phlebotomus) sergenti Parrot; Parrot, 1940 : 310.
Phlebotomus (Paraphlebotomus) sergenti Parrot; Theodor, 1948 : 97; 1958 : 21; Lewis, 1967 : 17; Perfil’ev,
1968 : 236; Artemiev, 1974a: 159, 160.
Perfil’ev and Theodor have given recent descriptions. The male has a slender basal lobe on the
coxite and four spines on a short style. The female has strong pharyngeal teeth and four or five
segments in the spermatheca.
2 (extra facts). Labrum and palpal sensilla as in P. papatasi. Hypopharynx with 16 teeth on each side.
Maxilla with seven lateral and 15 ventral teeth and a dental depth of 0-09 mm.
MATERIAL EXAMINED.
Pakistan: Ahmed Khel, 1 9.
DISTRIBUTION. India: Delhi, Karnal, Patiala (BMNH); Aurangabad (rare, Farooq & Qutubuddin,
1945 : 85); Banihal, Islamabad area, Srinagar (Jacob & Kalra, 1951 : 324); Abu Mount, Sambhar
(Jaswant Singh, 1933); Jaipur (Kaul et al., 1973 : 532); Mandi, Mendhar, Nowshera, Punch, Rajouri,
Riasi (Mitra, 1959 : 62); Maini Majera (Sinton, 19275 : 942); Agra, Aligarh (Uttar Pradesh), Ferozepore,
Karnal, Mohindergarh (Hariana), Narnaul, Rajkot, Roorkee (Sinton’s notes). Pakistan: Cherat, Chilas,
Dehra Ismail Khan, Gilgit, Gol, Gwadi, Jhelum, Keris, Lahore, Landi Kotal, Mir Muhammad, Parkuta,
Peshawar, Quetta, Rawalpindi, Said Pur, Shikarpur, Sukkur, Tank, Taxla (Lewis, 1967 : 17).
P. sergenti in Indo-Pakistan seemed to be confined to the plains north and west of the Bombay-—
Simla line (Sinton, 1932a), but extends a little beyond this area.
Subgenus SYNPHLEBOTOMUS Theodor
Phlebotomus subgenus Synphlebotomus Theodor, 1948 :97; 1958:22; Lewis & Ledger, 1976: 406.
Type-species: Phlebotomus martini Parrot, 1936, by original designation.
The hairy lobe of the coxite is medium-sized to large and the style has five spines. All but one
species of this small subgenus occur in tropical Africa.
236
Phlebotomus (Synphlebotomus) eleanorae Sinton
(Map 3)
Phlebotomus eleanorae Sinton, 1931a : 817; 1933e : 418. Holotype 3, INDIA (BMNH) [examined].
Phlebotomus (Phlebotomus) eleanorae Sinton; Parrot, 1940 : 310; 1946 : 67.
Phlebotomus (Synphlebotomus) eleanorae Sinton; Mesghali, 1965 : 267 [9]; Lewis & Ledger, 1976 : 406.
This is the only Oriental species in which the coxite is not very long and bears a distinctly visible
lobe, and the style is not very long and has five spines. The pharynx of the female has median
coarse teeth and the spermatheca 11—12 segments.
2 (extra facts). Labrum 0:25 mm long, 0-13 length of wing (1-94 mm), with wide upper brush-hairs in
short row. Hypopharynx with 16 teeth on each side. Antenna 3 = 0:19 mm long, 0-10 length of wing,
1-00 length of 4+ 5, 0-78 length of labrum.
MATERIAL EXAMINED.
Iran (Minab area): Chelo, 3.x.1964 (A. Mesghali), 1 9.
DISTRIBUTION. India: Karnal (Sinton, 1931a@: 817).
Subgenus LARROUSSIUS Nitzulescu
Phlebotomus subgenus Larroussius Nitzulescu, 1931 :274; Theodor, 1948:97; 1958:22; Perfil’ev,
1968 : 250. Type-species: Phlebotomus major Annandale, 19106, by original designation.
The coxite has no lobe, the style bears five spines, the paramere is not truncated and bears no
ventral process, and the spermatheca has an end-process. Two of the three Oriental species occur
in the west.
Study of the fascicle has shown that some species have more than four labral adoral sensilla,
and some have few maxillary lateral teeth.
Phlebotomus (Larroussius) betisi Lewis & Wharton
(Fig. 6, Map 3)
Phlebotomus (Larroussius) betisi Lewis & Wharton, 1963 : 117. Holotype 9, West MALAYsIA (BMNH)
[examined ].
The pharyngeal spicules are almost invisible, and the spermatheca has about 22 long bead-like
segments.
2 (extra facts). Labrum 0-35 mm long, 0-15 length of wing, with four adoral sensilla near the subapicals.
Pharynx with ridges and minute, almost invisible, spicules. Hypopharynx with 17 teeth on each side.
Antenna 3 = 3 mm long, 0-16 length of wing, 1-32 length of 4+5, 1-01 length of labrum, segment 5
without papilla. Maxilla with seven lateral and 21 ventral teeth and a dental depth of 0-11 mm; palpal
segment 3 with most sensilla near middle. Wing (2:23 mm) 3-2 times width, R./R,+. 3-60, R, overlap/R,
0-18.
MATERIAL EXAMINED.
West Malaysia: Betis, 1 9.
DIsTRIBUTION. West Malaysia: Betis, Kuala Trengan area (Lewis & Wharton, 1963).
Phlebotomus (Larroussius) kandelakii Shchurenkova
Phlebotomus kandelakii Shchurenkova, 1929 : 693. Syntypes 2 ¢, U.S.S.R. (found in Tbilsi Tropical
Institute) [not examined ].
Phlebotomus (Larroussius) kandelakii Shchurenkova; Theodor, 1958 : 23; Perfil’ev, 1968 : 261.
The pointed aedeagus has ventral teeth and the spermatheca is very long with 30-35 segments.
231
Phlebotomus (Larroussius) kandelakii burneyi Lewis
(Fig. 7, Map 3)
Phlebotomus (Larroussius) kandelakii burneyi Lewis, 1967:17; Artemiev, 1974a:160. Holotype 3,
PAKISTAN (BMNH) [examined].
P.k. burneyi differs from the nominate form in having two ascoids on antenna 3-7 in the male,
and other features.
2 (extra facts). Hypopharynx with 17 teeth on each side. Maxilla with four lateral and 17 ventral teeth
and a dental depth of 0-08 mm.
MATERIAL EXAMINED.
Pakistan: Gwadi, 1 9.
DISTRIBUTION. Pakistan: Gwadi, Kalam, Keris (Lewis, 1967).
Phlebotomus (Larroussius) keshishiani Shchurenkova
(Fig. 8, Map 4)
Phlebotomus keshishiani Shchurenkova, 1936 : 892. Syntypes 2? 3, U.S.S.R. (found in Tropical Institute of
Tadzhikistan (SSR), Dushanbe (= Stalingrad)) [not examined].
Phlebotomus (Larroussius) keshishiani Shchurerkova; Lewis, 1967 : 19; Perfil’ev, 1968 : 274.
P. keshishiani differs from P. major in having much longer genital filaments, the aedeagus very
narrow distally, and more spermathecal segments.
MATERIAL EXAMINED.
Pakistan: Parkuta, 1 9.
DISTRIBUTION. Pakistan: Gilgit, Parkuta, Rawalpindi, Said Pur (Lewis, 1967).
Phlebotomus (Larroussius) major Annandale
Phlebotomus major Annandale, 19105 : 46.
The aedeagus is long and narrow with nearly parallel sides, about as long as the paramere. The
species extends from the Mediterranean to Central Asia and northern India.
Phlebotomus (Larroussius) major major Annandale
(Fig. 9, Map 4)
Phlebotomus major Annandale, 1910 : 46; 1911c : 320; Newstead & Sinton, 1921 : 105; Sinton 1924a:
814; 1925d:107; 1927c: 948 [variation]; 1927d: 27; 1928c: 303 [synonymy]; 1932a:59; 1933e:
418; Sinton & Barraud, 1928 : 329. Lectotype 3, INDIA (Zoological Survey of India), designated by
Quate, 1962c : 157 [not examined].
Phlebotomus major var. grisea Annandale, 1911c: 320. Syntypes, sex not stated, INDIA (depository
unknown) [not examined]. [Synonymized by Quate, 1962c : 157.]
Phlebotomus major var. griseus Annandale; Sinton, 1932a : 59 [name emended; dark variant].
Phlebotomus (Phlebotomus) major Annandale; Parrot, 1940 : 310; Quate, 1962c : 157.
Phlebotomus (Larroussius) major Annandale; Theodor, 1958 : 250; Theodor & Mesghali, 1964 : 281;
Lewis, 1967 : 21; Perfil’ev, 1968 : 253.
The male of the nominate subspecies has a palpal formula of 1, 4, (2, 3), 5, and differs in a few
other respects from two Palaearctic subspecies in which the formula is 1, 4, 2, 3, 5. P. major major
was described in recent years by Perfil’ev and Theodor. It occurs in Pakistan and northern India.
In a male from Kasauli with an abnormal style one of the middle spines is replaced by two
narrow ones, and in a male from Sabadu one style has only one terminal spine.
238
2 (extra facts). Hypopharynx with 21 teeth on each side. Maxilla with four lateral and 26 ventral teeth,
dental depth 0-12 mm.
MATERIAL EXAMINED.
India: Dalhousie, 1 9.
DISTRIBUTION. India: Paresnath Hill (Annandale, 1912: 41); Bhowali, Dalhousie, vii.1906, Kasauli,
25.vi.1905, Simla (BMNH); Chamoli area (V. Dhanda); Banihal, Baramula, Islamabad, Jammu, Ramban
(Jacob & Kalra, 1951 : 324, 325); Mandi, Mendhar, Punch, Rajouri, Riasi (Mitra, 1959 : 59, 62); Kurse-
ong, Naini Tal (Quate, 1962c: 157); Chamba, Dehra Dun, Mukteswar (U.P.), Ranikhet, Sabathu
(Punjab) (Sinton’s notes). Nepal: Syabrudens (L. W. Quate, 28.x, 2.xi.1965, light trap). Pakistan: Ab-
bottabad, Rawalpindi, Said Pur (Lewis, 1967 : 21).
Early records have been checked because some (Sinton, 1927b) could have referred to P. longi-
ductus (=‘P. chinensis’) which was often found in the same sites (Sinton, 1928c : 306). P. m. major
seemed to occur all along the Himalayan foothills between 1555 and 2135 m, being, in India,
essentially a species of hills with marked summer rains (Sinton, 1932a).
Subgenus ADLERIUS Nitzulescu
Phlebotomus subgenus Adlerius Nitzulescu, 1931 : 275; Theodor, 1948 : 98; 1958 : 27; Perfil’ev, 1968 : 280.
Type-species: Phlebotomus chinensis Newstead, 1916, by original designation.
The coxite has no lobe, the style bears five spines, and the paramere is not truncated and carries
no ventral process. The spermatheca is incompletely segmented. In the male of most forms the
aedeagus has a subterminal minute fin-like barb. The subgenus is closely related to Larroussius,
and Parrot (1940) suggested possibly uniting them in view of their antennal formulae. The species
occur in temperate or in arid parts of the Old World, and two exist in the north of the Orient.
M. M. Artemiev (1977, in letter) is revising the Ad/erius of Afghanistan.
Phlebotomus (Adlerius) chinensis Newstead
Phlebotomus major var. chinensis Newstead, 1916 : 191.
The barb on the aedeagus is 10-35 um from the tip. The species was divided into several sub-
species but is here provisionally treated as comprising only chinensis, arabicus Theodor and
balcanicus Theodor, according to Artemiev’s tentative suggestions (see under P. /ongiductus).
Perfil’ev and Theodor gave recent descriptions. The species is almost entirely Palaearctic.
Phlebotomus (Adlerius) chinensis chinensis Newstead
(Map 4)
Phlebotomus major var. chinensis Newstead, 1916 : 191. LECTOTYPE 3, Cuina: ‘Wo Fu Hsu Temple,
1-6.vii.1914, co-type’ (BMNH), here designated [examined].
Phlebotomus chinensis Newstead; Sinton, 1928c : 306 [in part, synonymy]; 1932a:59; 1933d: 418;
Yao & Wu, 19415: 78.
Phlebotomus (Adlerius) chinensis Newstead; Theodor, 1958 : 28; Perfil’ev, 1968 : 280.
In the nominate form the barb of the aedeagus is 30-35 um from the tip (not more than 20 in
other forms) and the coxite brush has about 20 not very thick-standing hairs. P. c. chinensis is
one of the three Palaearctic taxa. Perfil’ev and Theodor gave recent descriptions.
2 (extra facts). Hypopharynx with 19 teeth on each side. Maxilla with four lateral and 19 ventral teeth
and a dental depth of 0:11 mm.
MATERIAL EXAMINED.
China: 2 2, 3 3 paralectotypes (BMNH), one labelled as type; same data as lectotype.
DIsTRIBUTION. China: Hainan (subsp.?, Leng, 1964 : 127); Kumming (subsp.?, Yao & Wu, 19416: 79).
239
Phlebotomus (Adlerius) longiductus Parrot
(Map 4)
[Phlebotomus chinensis; Sinton, 1928c : 306 [in part].]
Phlebotomus chinensis var. longiductus Parrot, 1928 : 29; 1940 : 310 [¢ ascoid formula variable]; 1946 : 68.
Syntypes 2 g, U.S.S.R. (depository unknown) [not examined].
Phlebotomus (Adlerius) chinensis hindustanicus Theodor, 1958: 29, 30. Syntypes 3, NORTH-WEST OF
INDIAN SUBCONTINENT (TC) [not examined]. [Synonymized by Lewis, 1967 : 21.]
Phlebotomus (Adlerius) chinensis longiductus Parrot; Theodor, 1958 : 29; Theodor & Mesghali, 1964 : 193
[?]; Lewis, 1967 : 21 [¢ ascoid formula variable]; Perfil’ev, 1968 : 285.
Phlebotomus (Adlerius) longiductus Parrot; Artemiev, 1974a : 163.
In the male the barb of the aedeagus is shallow and about 12-14 um from the tip, and the coxite
brush has 50-60 hairs.
Artemiev treated this form as a species in view of differences from Chinese P. chinensis in the
number of hairs on the coxite and the shape and position of the aedeagus barb, sympatric relation
to form halepensis Theodor and P. simici Nitzulescu, and sharp differences from the allopatric
‘subspecies’ of P. chinensis except arabicus Theodor, balcanicus Theodor and tauriae Perfil’ev,
1966 : 312, which could be subspecies of P. Jongiductus. He pointed out that P. Jongiductus occurs
in southern U.S.S.R., Iran, Afghanistan and northern Pakistan, and that in Afghanistan it is a
mountain cold-resistant species occurring from 1000 to 2800 m.
Recent descriptions have been given by Artemiev, Lewis, Perfil’ev and Theodor. In an occasional
male from India one of the ascoids on antenna 8 is vestigial.
2 (extra facts). Labrum 0-40 mm long, 0-14 length of wing (2°76 mm). Hypopharynx with 21 teeth on each
side. Maxilla with three lateral and 23 ventral teeth and a dental depth of 0-12 mm.
MATERIAL EXAMINED.
Pakistan: Gwadi and Keris, 2 9.
DIsTRIBUTION. India: Bhowali, Kasauli, Simla (BMNH); Banihal (Jacob & Kalra, 1951 : 325); Punch
(Lewis, 1967 : 23); Mandi, Mendhar (Mitra, 1959 : 62); Kurseong, Ranikhet (Sinton’s notes). Nepal:
Chobhar (Jane Wilson, 1976, in cave); Syabrudens (L. W. Quate via BPBM, 1965). Pakistan: Gilgit, Gol,
Gwadi, Keris, Parkuta, Said Pur (Lewis, 1967 : 23).
In India P. longiductus was not usually found below 1555 m and occurred in the same area as
P. major, according to Sinton (1928c; 1932a).
Subgenus EUPHLEBOTOMUS Theodor
Phlebotomus subgenus Euphlebotomus Theodor, 1948:98; 1958:32; Hennig, 1972:53 [related to
Phlebotomus and Anaphlebotomus]. Type-species: Phlebotomus argentipes Annandale & Brunetti, 1908,
by original designation.
The coxite has no lobe, the style bears five spines, and the paramere has three processes. Hennig’s
(1972) study of the paramere of a fossil species suggests that this is an ancient subgenus. There
are five species in the Old World.
Phlebotomus (Euphlebotomus) argentipes Annandale & Brunetti
(Figs 10-16, Map 4)
Phlebotomus argentipes Annandale & Brunetti in Annandale, 1908 : 101 [authorship according to ICZN
Article 51(c)]; Annandale, 19106 : 42; 1911a: 159; 1911c : 309; Howlett, 1915 : 294; Franga, 1922 : 14;
Sinton, 1924a : 814; 1925c¢ : 789 [colour variation]; 1925d : 107; 1927d: 27; 1928c : 301 [synonymy];
1932a : 59; 1933d : 227; 1933e : 418; Christophers & Barraud, 1926 : 853 [?]; Christophers, Shortt &
Barraud, 1926 : 177; Shortt, Barraud & Craighead, 1926 : 330; Sinton & Barraud, 1928 : 329; Mukerji,
1931 : 441 [larva]; Raynal, 19355 : 245; Raynal & Gaschen, 1935g : 737 [figs in 1935/]; Keilin & Tate,
1937 : 254; Mitra, 1952 : 550 [palp sensilla]; 1956 : 229; Mitra, 1953 : 434; Mitra & Roy, 19536 : 369
240
[variation]. Lectotype 3, INDIA (Zoological Survey of India), designated by Quate, 1962c : 157 [not
examined ].
[Phlebotomus zeylanicus Annandale; Annandale, 1910a: 61. Misidentification according to Sinton,
1924a : 813; 1928c : 301.]
Phlebotomus marginatus Annandale, 1910a: 62; Sinton, 1932a:59 [colour form]. Holotype 2, Sri
LANKA (depository unknown) [not examined]. [Synonymized by Theodor, 1948 : 108.]
Phlebotomus argentipes var. marginatus Annandale; Annandale, 19115: 203; 1911c: 319.
Phlebotomus annandalei Sinton, 1923a : 744; 1924a : 815. Holotype ¢ (ICZN Article 73(a)), INDIA (deposi-
tory unknown) [not examined]. [Synonymized by Sinton, 192Sc : 789.]
Phlebotomus (Phlebotomus) argentipes Annandale & Brunetti; Parrot, 1937: 116; 1940: 310; 1946 : 68;
1953 : 114; Lewis, 1957: 165; Quate & Fairchild, 1961 : 211; Quate, 1962b : 254; 1962c : 157 [syno-
nymy ].
Phlebotomus (Euphlebotomus) argentipes Annandale & Brunetti; Theodor, 1948 : 99; Lewis, 1967 : 23;
1973a : 246; 1973c : 147; Lewis & Killick-Kendrick, 1973 : 4.
Phlebotomus argentipes var. glaucus Mitra & Roy, 1953 : 372. 10 9 syntypes, INDIA (Museum of the
Armed Forces Medical College, Poona) [not examined]. [Synonymized by Lewis, 1967 : 24.]
In the female the pharynx has a group of spines, and the spermatheca a large apical segment,
and in the male a pair of spines lies parallel to the aedeagus, the paramere has three lobes and
the style five spines.
Descriptions include those of Quate & Fairchild, Raynal (19355) and Sinton (1925c).
The form g/aucus is here treated as a synonym in the light of present knowledge of variation.
(East India, Calcutta, Howrah). Eye 0:56 length of head. Labrum 0-25 (0:23-0:27) mm long, 0-12
(0:12-0:13) length of wing, shoulders angular, most sensilla small, the two mid-apical ones prominent.
Cibarium with distinct spicules. Pharyngeal armature with median anterior teeth and lateral and posterior
spiculate ridges. Hypopharynx with about 16 long teeth on each side. Antenna 3 = 0-22 (0:20-0:23) mm
long, 0-11 (0:10-0:11) length of wing, 1-25 (1-:25—1-34) length of 4+ 5, 0-87 (0:80-0:92) length of labrum,
no papilla on 5, two ascoids on segments 3-15, that on 4 being 0-41 (0:30-0:53) length of segment.
Mandible with very fine teeth about 1:2 um wide. Maxilla with 8-8 (7-11) lateral teeth, 14-9 (13-17)
ventrals, and a dental depth of 0-08 mm; palpal ratio 10 : 22 : 31 : 16 : 34, clavate sensilla close together
around middle of segment 3. Scutum dark brown, pleuron pale, inter-precoxal lobes normal, mesane-
pisternum with about five to eight lower hairs. Wing length 2-06 (1:92-2:22) mm, 3-2 times width,
R,/Ro43 1:91 (1:59-2:06), R, overlap/R, 0:14 (0:09-0:20). Tarsi appearing silvery in some lights. Sper-
matheca carrot-shaped with about 15 segments, the end one large, outer wall of common duct diverging
distally.
3 (Calcutta area, extra facts). Eye 0:56 length of head. Style 0-62 length of coxite, 5-8 times as long as
thickness (basad of middle spines).
VARIATION. This species was known to show some geographical variation (Lewis, 1957), and
further observations were made on the limited material available owing to the importance of the
species and its biological variation. In parts of India it bites man readily, is peridomestic, and is
an important vector of kala-azar, whereas in south-east Asia it seldom if ever attacks man, and
in Sri Lanka may be strongly zoophilic (Lewis & Killick-Kendrick, 1973). It has even been
suggested, by R. S. Bray in 1974, that P. argentipes may be a species complex including a zoophilic
and an anthropophilic species in one area.
Available specimens were limited, but those from several areas were examined for comparison
with P. argentipes from the area around the type-locality of Calcutta, with the following results.
In a few flies from some areas one maxillary lateral tooth is vestigial and makes counting difficult.
South India in general. Labrum 0:24 (0:22-0:27) mm long, 0-12 (0:12-0:13) length of wing. Antenna 3 0-22
(0-19-0-23) mm long, 0:11 (0:10-0:12) length of wing, 1-21 (1-18-1-25) length of 4+5, 0-88 (0:84-0:93)
length of labrum. Ascoid on 4 = 0-49 (0:39-0:60) length of segment. Maxilla with 7-0 (6-11) lateral and
13-3 (14-19) ventral teeth. Wing length 2-03 (1:82-2:22) mm, R./Ro3 1:74 (1:42-1:95), R, overlap/R.
0-15 (0-09-0:21).
North India in general. Labrum 0:25 (0:24-0:27) mm long, 0-12 (0:11-0:12) length of wing. Antenna
3 = 0-22 (0:20-0:26) mm long, 0-10 (0:10-0:12) length of wing, 1:21 (1:14-1:33) length of 4+5, 0-88
(0-80-0-94) length of labrum. Ascoid on antenna 4 = 0-52 (0:33-0:74) length of segment (the highest
value is in one of two flies from Bombay, with an antenna 3/labrum value of 0:83; otherwise the mean is
241
Figs 10-18 Phlebotomus species. 10-16, P. argentipes: (10-12) 9°, heads from India (Howrah), Sri
Lanka (Pannipitiya) and West Malaysia (Lamir) (scales adjusted for wing length); (13) 9, sperma-
theca (Venkatapuram); (14, 15) base of its duct (Venkatapuram and Lamir); 16, 3, aedeagus and
parameres (Lamir). 17, P. kiangsuensis, 2, labrocibarium. 18, P. philippinensis gouldi, 2, spermatheca.
242
0:50 and the maximum 0-64). Maxilla with 8-5 (8-9) lateral and 15-0 (12-18) ventral teeth. Wing length
2:13 (2:01-2:37) mm, R2/Re+3 1°77 (1:51-2:25), R, overlap/R, 0-12 (0:00-0:18). In a female from Katihar
one (short) ascoid on antennal segment 3 has a bifid tip, a feature seen in some ascoids of a few Calcutta
flies (S. Das, 1975, in litt.). In two females, outside the series, from Hosur (H. Trapido), the ascoid on 4 is
0-39 and 0-70 of the segment length.
India: various. (1) Northern hills. Labrum 0-25 (0:23-0:27) mm, 0-11 (0:10-0:12) length of wing.
Antenna 3 = 0:23 (0:21-0:25) mm long, 0:10 (0:10-0:12) length of wing, 1:29 (1:24-1-36) length of
445, 0-93 (0:85-1:01) length of labrum, ascoid on 4 = 0-65 (0:64-0:67) length of segment. Maxilla with
10-3 (9-11) and 16-3 (14~18) lateral teeth. Wing length 2-21 (2:08-2:32) mm, R./R2,3 1°71 (1:52-1-80),
R, overlap/R, 0-10 (0:04-0:11). (2) Near the hills of Kerala. In the single female the labrum is 0-14 as long
as the wing, the ascoid on antenna 4 is 0-74 of its length, the maxilla has 12 lateral and 16 ventral teeth,
and the wing length is 2:22 mm. The ascoid alone was examined in six flies from Assam (Gologhat) and
is 0-62 (0:53-0:67) the length of segment 4. In one female from Mount Abu the figure is 0-46. In these
rather peripheral areas the ascoid tends to be long. The ascoid ratio in a fly from Nepal is 0-60.
Sri Lanka. Labrum 0:24 (0:22-0:26) mm long, 0-12 (0:12-0:12) length of wing. Antenna 3 = 0:22
(0:19-0:24) mm long, 0-11 (0:10-0:11) length of wing, 1-24 (1-19-1-31) length of 4+5, 0-90 (0:83-0:94)
length of labrum, ascoid on 4 = 0-75 (0:68-0:86) length of segment. Maxilla with 8-9 (8-10) lateral and
13-8 (11-17) ventral teeth. Wing length 2-00 (1:86-2:13) mm, R,/R.+3 2:00 (1:70-2:31), R, overlap/R,
0-18 (0:14-0:26). The mean ascoid length, R,/R,.3; and R, overlap are high. Females are distinguishable
from all the east India and south-India-general flies, and all but one of those from the general area of
north India.
Thailand. Labrum 0:24 (0:23-0:26) mm long, 0:11 (0-11-0-11) length of wing. Antenna 3 = 0:25
(0:23-0:29) mm long, 0-11 (0-10-0-12) length of wing, 1-12 (1:07-1:20) length of 4+5, 1-04 (0-95-1-08)
length of labrum, ascoid on 4 = 0-67 (0:57-0:72) length of segment. Maxilla with 8-7 (8-9) lateral and
12:2 (11-13) ventral teeth. Wing length 2-27 (2:13-2:45) mm, R,/R2,3 2°12 (1:97-2:65), R, overlap/Re2
0-11 (0-07-0-18). The long antenna 3, ascoid and R, are close to West Malaysian values, and the wing
length is rather high and the number of maxillary ventral teeth low. In South Vietnam the ascoid is rather
long (Raynal, 19355).
West Malaysia. Labrum 0:23 (0:22-0:24) mm long, 0:11 (0:10-0:11) length of wing. Antenna 3 = 0-24
(0:22-0:25) mm long, 0-11 (0-10-0-12) length of wing, 1:14 (1:11-1:17) length of 4+5, 1-05 (0-95-1-10)
length of labrum, ascoid on 4 = 0-77 (0-69-0-81) length of segment. Maxilla with 8-1 (6-11) lateral and
13-8 (11-17) ventral teeth. Wing length 1-96 (1:74-2:26) mm, R,/R2.3 1:96 (1:74-2:26), R, overlap/R2
0:16 (0:09-0:24). The labrum is rather short, antenna 3/4+5 low, antenna 3/labrum value high, ascoids
long, and R,/R.,3 and R, overlap /R, values high. All are distinguishable from east India, south-India-
general and north-India-general flies; the only one of the latter with a long ascoid has a short antenna
3 and a low antenna 3/labrum value. The West Malaysian form, like the Sri Lanka form, has a long ascoid,
but is distinguished from it by a lower labrum/wing length value, a lower antenna 3/4+5 value, and a
higher antenna 3/labrum value. In the male of both east India and West Malaysian forms the paired
ascoids end at antennal segment 10.
Over-all minima and maxima. The data from all areas give the following result; labrum 0:22-0:27 mm
long, 0:10-0:14 length of wing; antenna 3 = 0:19-0:29 mm long, 0-10-0-12 length of wing, 1:11-1:36
length of 4+5, 0-80-1-10 length of labrum; ascoid on 4 = 0-30-0-86 length of segment; maxilla with
6-12 lateral and 11-19 ventral teeth; wing length 1:82-2:45 mm, R,/R2,3 1:42-2-65. R, overlap/R,
0-00-0-26.
COMMENTS. The specimens examined show similarities between flies from east India and the
general areas of south and north India, and they indicate Sri Lanka and West Malaysian forms
(with long ascoids and other features) distinct from these. Some specimens from the peripheral
areas of India and from north-east Thailand gave intermediate measurements, the differences
are small, and little is known of the species in Burma and Thailand. Therefore none of the
variants is treated here as a subspecies. There is, however, a striking morphological difference
between the short ascoid of the eastern Indian vector of kala-azar and the long ascoid of the
non-anthropophilic P. argentipes of south-east Asia (Figs 10, 12).
There is nothing to suggest the existence of two species in the P. argentipes complex in India,
apart from the finding of long and short ascoids at Bombay and at Hosur, but the lack of man-
biting P. argentipes in the Sagar (Shimoga) area (noted under distribution below) is interesting.
No P. argentipes were found in a very large collection from Perak, and there may be some dis-
continuity of distribution which would suggest a subspecific status for some variants.
243
Further study of this important species would be instructive, and could well include cyto-
taxonomy and measurement of more specimens from a wider area, and perhaps the relation of
leg measurements to wing length.
MATERIAL EXAMINED.
India. East: Howrah (Calcutta, S. Das), 10 2; north, general: Aurangabad, Chindwara, Jalna, Karnal,
Parel (Bombay, 2), Ranchi, Saharanpur, Sambhar, 10 2; south, general: Puligumma, Undi (3), Venkata-
puram (2) 6 2; various, northern hills: Kathgodam, Simla (3), 4 9; various, other: Gologhat, 6 2; Kula-
thupurzha, 1 2; Mount Abu, 1 9. Nepal: Dhunibesi (21.vi.1961, Y. Shogaki, 1 2). Sri Lanka: Pannipitiya
(18.iv.1973, R. Killick-Kendrick, 10 ¢). Thailand: Ban Bon Dan (11.xii.1975, D. J. Gould, 6 2). West
Malaysia: Lamir, 10 9.
DISTRIBUTION. Bangladesh: Dhurmakura (Sinton’s notes). Borneo (Sabah): Bum Bum Island (Lewis,
1968: 11); Kalabakan area (Quate & Fairchild, 1961 : 213; Quate & Rosario, 1962: 791). Burma:
Rangoon (probably, Sinton, 1928c : 311; 1932a: 70). India: Howrah (as above); Katiher, Kotelanka,
Kulathurpuzha, Mount Abu, Nagpur, Nalbari, Ranchi, Simla (BMNH); Calcutta (Basu & Ghosh,
1954a : 1955); Aurangabad, Jalna, Patan (Farook & Qutubuddin, 1951 : 85); Kathgodam (14.x.1907, on
pony, J. D. E. Holmes); Ajmer, Sambhar (Jaswant Singh, 1953); Hoora (Mitra, 1953 : 158); Khandwa,
Kirki, Mahableshwar, Mahad, Panchgani, Pashan, Poladpur, Wai etc. (Mitra, 1954b: 111; 1955 : 82);
Hyderabad (near lake, not numerous, Qutubuddin, 1944 : 208); Chamoli area (Rao et al., 1973); Panada
Agraharam, Undi, Vellore (6 3 on bullock), Vekatapuram (R. Reuben); Asansol Kamptee, Palod, Poona,
Port Canning (Sinton, 1924 f: 1041); Golaghat, Lucknow, Puri, Purneah, Rajmahal (Sinton, 1925c : 789);
Faizabad, Pamban Island, Patna, Sarahanpur (Sinton, 1927b: 942); Sanawar (Sinton, 1932a: 70);
Gauhati, Karnal, Narnaul, Nedumangad, Parbatipur, Rajbari, Trivandrum (Sinton’s notes); Sagar
(Shimoga) area (Hosur, Kannur, Konehosur, Kumsi, H. Trapido; Work et al., 1957; Trapido ez al.,
1959; no phlebotomines found biting); Bombay (first record, widespread, Young, 1927 : 679). Indonesia:
Denpasar area (Lewis & Dyce, 1976 : 208). Laos: Luang Prabang (Quate, 1962b : 256). Nepal: Kathmandu
area (as above). Pakistan: Lahore, Mir Muhammad, Taxla (Lewis, 1967 : 24). Sri Lanka: (not plentiful,
Smith, 1959: 17); Balangoda area (Jun Akiyama, 12.viii.1977, 4 9, 51 3 from cow-baited trap net at
Kaltota); Peradeniya (Annandale, 1910a : 59; 19116 : 203; 1911c : 319); Delft Island (Carter & Antoni-
pulle, 1949 : 68); Depanama, Kalagoda, Pannipitiya (Theodor, 1938a : 269). Thailand: Ban Bon Dan,
Khao Yai (as above); Chieng Mai (Quate, 19625 : 256). Vietnam (South): 22 km south of Nha Trang
(Quate, 19625 : 256); Duc Pho (Raynal, 1936a : 360). West Malaysia: Kuantan area (box traps near
Lamir), Rantan Panjang, Ulu Gombak (Lewis, 1957: 106); Gua ‘Che Yatim (Lewis & Wharton,
1963 : 118); Batu Caves (Quate, 1962a : 226; Quate & Fairchild, 1961 : 212).
Sinton (1932a) considered that in India P. argentipes occurred in a moist climate mainly east and
south of the Bombay-Simla line, but with a focus in the Kathiawar Peninsula. Sanawar, at
1220 m, was unusually high for it (Sinton, 1927g), but Mitra (1954a) found it at nearly 1555 m
in the Bombay area. Smith’s (1959: 17) remarks on patchy distribution in India may explain
diverse findings in south-east Asia. In 1975 P. argentipes was reported from Iran.
Phlebotomus (Euphlebotomus) kiangsuensis Yao & Wu
(Fig. 17, Map 4)
Phlebotomus sp. Raynal, 1937 : 83 [China; microfilariae in midgut].
Phlebotomus kiangsuensis Yao & Wu, 1938 : 527; 1941b : 78. Holotype 3, CHINA (depository unknown)
[not examined]. Conditional name available under ICZN Article 17(8).
Phlebotomus (Euphlebotomus) kiangsuensis Yao & Wu; Theodor, 1948 : 108; 1958 : 32.
Phlebotomus (Phlebotomus) kiangsuensis Yao & Wu; Lewis & Wharton, 1963 : 120 [variation]; Cates &
Lien, 1970 : 538.
The male differs from that of P. argentipes in having a short aedeagus and the central lobe of
the paramere thicker than the upper lobe. P. kiangsuensis, first found in the Palaearctic Region,
has been redescribed by Theodor (1958).
? (China, extra facts). Hypopharynx with about 18 teeth on each side. Antenna 5 with papilla. Maxilla
with eight lateral and 21 ventral teeth, and a dental depth of 0-11 mm.
3 (China, extra facts). Antenna 4 apparently with one ascoid, antenna 5 bearing one normal ascoid,
one very small one, and apparently a papilla.
244
VARIATION. Males from West Malaysia show reduction of proximal ascoids (Lewis & Wharton,
1963) and apparently no papilla on antenna 5. Further study of these delicate structures is
desirable.
MATERIAL EXAMINED.
China: Kiangsu (3 9, 1 ¢ bred in laboratory, in BMNH collection, evidently originally treated with strong
potash which makes papillae difficult to see). West Malaysia: 4 2, 3 3.
DISTRIBUTION. China: Kukong area (Chen & Hsu, 1955 : 302). Taiwan: Taitung area (3 2, Cates & Lien,
1970 : 539). West Malaysia: Batu Cave (5.ix.1959, N. E. McClure); Betis (Lewis & Wharton, 1963 : 120).
P. kiangsuensis and two other species of Euphlebotomus occur in Palaearctic China.
Phlebotomus (Euphlebotomus) philippinensis Manalang
Phlebotomus philippinensis Manalang, 19306 : 175.
The male differs from that of P. argentipes in having longer eyes (as in the female) and a deeper
paramere, and in the relative lengths of the main and middle lobes of the paramere.
Quate & Rosario (1962) gave a recent description and indicated that sympatric females of the
two species would be difficult to separate and that records should be confirmed by means of
males; also that P. philippinensis is smaller, and has less developed cibarial spicules, a stronger
chitinous arch and slightly longer antenna 3.
Phlebotomus (Euphlebotomus) philippinensis gouldi Lewis subsp. n.
(Figs 18-21, Map 4)
P. p. gouldi has higher antenna 3/wing length and antenna 3/4+5 values in both sexes than does
P. p. philippinensis, antenna 3 in the female reaches beyond the labrum, the maxillary ventral
tooth number and dental depth are greater, wing length is greater in the male, R, overlap is
longer in both sexes, and the style is rather long and narrow.
9. Eye 0-43 length of head. Labrum 0-22 (0:22-0:23) mm long, 0-11 (0:11-0:11) length of wing. Antenna 3
extending beyond labrum, 0-34 (0:31-0:36) mm long, 0-17 (0-17-0-18) length of wing, 1-24 (1-:23-1-26)
length of 4+5, 1-42 (1:39-1-45) length of labrum, two ascoids on segments 3-15, that on 4 = 0-74
length of segment, papilla on 5. Maxilla with seven lateral and 12 ventral teeth and a dental depth of
0-06 mm, palpal ratio 10: 24: 34: 19: 36. Wing length 1-98 (1:91-2:05) mm, 3-1 times width, R./R».5
1:77 (1-:70-1-84), R, overlap/R, 0-13 (0:08-0:17). Spermathecae delicate and indistinctly wrinkled or
segmented, with large end-segment, individual ducts delicate and common duct thick-walled.
3. Eye 0-42 length of head. Labrum 0:20 mm long, 0-10 length of wing. Antenna 3 = 0-40 mm long,
0-20 length of wing, 1:21 length of 4+5, 2-02 length of labrum; two ascoids on segments 3-13, one on
14 and 15, that on 4 = 0-58 length of segment. Wing length 1-95 mm, 3-4 times width, R,/R,,3 1-58, Ry;
overlap/R, 0-13. Style 0-61 length of coxite, 3-6 times as long as width (basad of middle spines).
MATERIAL EXAMINED.
Holotype 2, Thailand: Ban Bon Dan, 11.xii.1975 (D. J. Gould), light trap with CO, in evergreen and
deciduous tropical forest (BMNH).
Paratypes. Same data, 10 and 11.xii.1975, 1 9, 1 ¢ (BMNH).
DISTRIBUTION. Thailand: Ban Bon Dan (as above); Khao Yai (25.iii.1976, D. J. Gould, 700m, rain
forest, 1 9).
Phlebotomus (Euphlebotomus) philippinensis philippinensis Manalang
(Map 4)
Phlebotomus philippinensis Manalang, 1930b : 175; Sinton, 1931d: 104. Syntypes 2 3, PHILIPPINES (des-
troyed according to Quate & Rosario, 1962 : 789) [not examined].
Phlebotomus (Euphlebotomus) philippinensis Manalang; Theodor, 1948 : 108.
Phlebotomus (Phlebotomus) philippinensis Manalang; Quate, 1965 : 20; Quate & Rosario, 1962 : 789.
245
——|
705
Figs 19-26 Phlebotomus species. 19-21, P. philippinensis gouldi, 3, terminalia, aedeagus and para-
meres, and tip of paramere. 22-26, P. hoepplii, after Tang & Maa: (22) 9, pharynx; (23) ?, sperma-
theca; (24) 3, pharynx; (25) 3, terminalia; (26) 3, aedeagus and parameres.
246
2 (extra facts). Eye 0:38 length of head. Labrum 0-23 mm long, 0-13 length of wing. Hypopharynx with
16 teeth on each side. Antenna 3 not reaching tip of labrum, 0:24 mm long, 0-13 length of wing, 1-15
length of 4+5, 1-04 length of labrum, ascoid on 4 about 0-8 length of segment, papilla present on 5.
Maxilla with seven lateral and 17 ventral teeth and a dental depth of 0:20 mm. Wing length 1-81 mm,
R,/Re+3 1°77, Ry apex/R, 0-22. Spermatheca with faint pit and knob; common duct with thick walls.
3 (extra facts). Eye 0-41 length of head. Labrum 0-16 (0:15-0:16) mm long, 0-09 (0:09-0:09) length of
wing. Antenna 3 = 0:26 (0:24-0:28) mm long, 0-16 (0-15-0-16) length of wing, 1-16 (1:10-1:22) length
of 4+5, 1-67 (1:63-1:71) length of labrum, ascoid on segment 4 = 0-74 length of segment, papilla
present on 5. Wing length 1-66 (1:55—1-76) mm, R,/R2+3 1°62 (1:38-1-86), R, overlap/R, 0:51 (0-08-1-04).
Lower lobe of paramere a broad flange extending mesally and bearing four or five spines. Style 0-54
length of coxite, 4-1 as long as width (basad of middle spines).
MATERIAL EXAMINED.
Philippines: Imus, 1 3; La Mesa, 1 9; Silang, 1 3.
DISTRIBUTION. Philippines: Novaliches (Manalang, 19305 : 175); Cotabato, Eran area, Los Arcos etc.
(Quate, 1965 : 20); Imus, La Mesa, Silang (Quate & Rosario, 1962 : 791).
Subgenus ANAPHLEBOTOMUS Theodor
Phlebotomus subgenus Anaphlebotomus Theodor, 1948 : 99; Hennig, 1972 : 53. Type-species: Phlebotomus
stantoni Newstead, 1914, by original designation.
The coxite has no lobe and the style bears four spines. It is represented in the Region by three
species which, between them, cover a wide area.
Phlebotomus (Anaphlebotomus) colabaensis Young & Chalam
(Map 5)
Phlebotomus colabaensis Young & Chalam, 1927 : 859; Sinton, 1932a : 59; 1933d: 226 [2]; 1933e: 418
[$]. Holotype 3, INDIA (Central Research Institute, Kasauli) [not examined].
Phlebotomus (Anaphlebotomus) colabaensis Young & Chalam; Theodor, 1948 : 108; Lewis, 1967 : 24.
The paramere is bilobed and the style has four spines. The following notes are based on Young
& Chalam and on Sinton, and study of one fly.
?. Labrum 0-30 mm long, 0-15 length of wing. Hypopharynx with about 19 teeth on each side. Maxilla
with 11 lateral and 23 ventral teeth and a dental depth of 0:11 mm. Antenna 3 = 0:29 mm long, 0:14
length of wing, 1-18 length of 4+ 5, 0-97 length of labrum, ascoid on segment 3 = 0-61 length of segment,
papilla on 5. Palpal segment 3 with sensilla grouped near middle. Thoracic sternal lobe broad. Sperma-
theca not markedly carrot-shaped, with small end-segment; duct long, about four times length of sper-
matheca, joining common duct.
3d. Antenna 3 = 0:22 mm long, 1:1 length of 4+5. Palpal formula 1-4—2-(3-5), ratio 10 : 27 : 40 : 20: 40.
Wing length 1-47 mm, width 0-41 mm, R,/R,,3 1:4, R, overlap/R, 0-1. Hind tibia 1-72 length of femur.
Aedeagus pyramidal and sharply pointed. Paramere bilobed, lower lobe bare and mammiform. Style
about 3-9 times as long as wide, with four spines of nearly equal thickness, at 0-23, 0:57, 0-73 and tip.
MATERIAL EXAMINED.
India: Kulathurpuzha, 1 2.
DISTRIBUTION. India: Kulathurpuzha (BMNH); Sambhar (Jaswant Singh, 1933); Hyderabad (near river,
Qutubuddin, 1944 : 208); Bombay (Sinton, 1928c : 310; Young & Chalam, 1927 : 49); Bissamcuttack
(Sinton, 1932a : 71). Pakistan: Lahore (Lewis, 1967 : 24).
Phlebotomus (Anaphlebotomus) hoepplii Tang & Maa
(Figs 22-26, Map 5)
Phlebotomus hoepplii Tang & Maa, 1945 : 25, Holotype 3, CHINA (T. Maa’s collection) [not examined].
247
The male differs from that of P. stantoni in having the spine near the aedeagus much longer
than it.
The following description is adapted from the original for which five flies of each sex were used
for most measurements.
?. Pharynx about three times as long as wide, armature with faint transverse and slightly oblique lines;
anteriorly lines are more developed and there are lateral backward-pointing teeth and medially some
small ones. Antenna 3 = 0:25 (0:24-0:27) mm long, 1-2 (1:1—-1-3) length of 4+ 5, two ascoids on segments
3-15, that on 4 almost as long as segment and reaching beyond its tip. Palpal formula 1-4-2-3-5, ratio
10: 31:44:17: 34. Wing length 1-86 (1-75—2-07) mm, width 0-55 (0:48-0:61) mm, R,/R2+3 2:1 (1-8-2:3),
R, overlap/R, about 0:25. Hind tibia 1-84 (1:77-1:91) length of femur. Spermatheca fusiform with small
head and short neck; roughly 14 segments figured; duct shorter than spermatheca; common duct long.
$. Cibarium unarmed. Pharynx about four times as long as wide, armature with a series of oblique
ridges radiating from the middle, and antero-lateral sharp teeth pointing postero-medially. Antenna
3 = 0:27 (0:26-0:28) mm long, 1-26 (1:23-1:33) length of 4+ 5; two ascoids on segments 3-15, that on
4 about 0-83 length of segment and reacing its tip. Wing length 1-68 (1:58-1:75) mm, width 0-50
(0:46-0:53) mm. Hind tibia 0-20 length of leg. Aedeagus with rounded tip, and a long spine on each side.
Paramere with three lobes, the small ventral one usually bearing three curved spines. Coxite long and
slender with a large group of long hairs. Style spindle-shaped with four spines, the two proximal ones
being more slender, at 0-48, 0-53, 0-91 and tip. Cerci narrow and club-like distally, not figured.
DISTRIBUTION. China: Kuang-chou (Chen & Hsu, 1955 : 302); Masha, Shao-wu (Tang & Maa, 1945 : 25).
Phlebotomus (Anaphlebotomus) stantoni Newstead
(Figs 27-29, Map 5)
Phlebotomus stantoni Newstead, 1914 : 190; Sinton, 1923a : 749; 1928c : 311; 1931d: 99, 104; 1933d: 226;
Raynal & Gaschen, 1934d: 670 [3]; Raynal, 19355 : 237 [3]; 1936a : 357; Theodor, 1938 : 269; Yao
& Wu, 1940: 773; 19416: 74; Tang & Maa, 1945: 29. Holotype 2, West MALAysIA (BMNH) [ex-
amined ].
Phlebotomus (Phlebotomus) stantoni Newstead; Parrot, 1946 : 67; Parrot & Clastrier, 1952 : 154; Quate &
Fairchild, 1961 : 205; Quate, 1962 : 256.
Phlebotomus (Anaphlebotomus) stantoni Newstead; Theodor, 1948 : 99.
Phlebotomus maynei Sinton, 1930b: 195. Holotype 3, INDIA (BMNH) [examined]. [Synonymized by
Raynal & Gaschen, 1934d : 670.]
Phlebotomus (Phlebotomus) maynei Sinton; Parrot, 1940 : 310; Lewis, 1967 : 27; 19745 : 190.
The female has individual spermathecal ducts longer than the spermathecae and a very long
thick-walled common duct. In the male the rods near the aedeagus are not longer than it. Most
of the following description is adapted from those of Newstead, Parrot & Clastrier and Sinton
(1923a; 193ld; 1933d), and based partly on the holotype. The species was figured by these
authors and by Quate.
Quate & Fairchild (1961) mention a pigment patch, but Raynal (19355), Raynal & Gaschen
(1931d), Sinton (1931d) and Yao & Wu (1940) state that there is none, and I have not found one.
. Labrum 0:23-0:26 mm long. Cibarium with about 15 pointed denticles of various lengths, irregularly
arranged, the two or three median ones longer and stouter than the others. Pharynx two or less times wider
posteriorly than anteriorly, with numerous posterior transverse folds, parallel and concentric, the hinder
ones bearing short fine spicules; before the folds are numerous long strong pointed spines in a convex
crescent. Hypopharynx with about 16 teeth on each side. Antenna 3 = 0:27-0:30 mm long, longer than
4+5, 1-1 length of labrum, two long ascoids on segments 3-15, those on 4 about 0-8 length of segment
and reaching to or beyond its tip. Maxilla with nine lateral and 18 ventral teeth and a dental depth of
0-08 mm. Palpal formula 1-4—2-3-5 or 1-4-2-(3-5), ratio 10:29:46:19:50. Wing length 1-9 mm,
width 0-58-0-61 mm, R,/ Re; 1-6-1:7, R, overlap 0:08-0:12 mm. Hind legs 3-55—3-75 mm long. Sperma-
theca fusiform, with 15 or 16 rings, neck thick and short, head more or less oblong, duct striated, common
duct very long, about 1-5 length of individual ones, with thick wall.
3. This was described mainly by Raynal & Gaschen (1935d) and Raynal (19350).
248
Figs 27-37 Phlebotomus species. 27-29, P. stantoni, 2 (from Java), cibarium and spermathecae.
30-37, P. teshi, 2: (30, 31) tips of labrum and hypopharynx; (32) cibarium; (33) antennal segments
4, 15 and 16; (34) mandible and maxilla; (35) two hairs, a scale and a sensillum from palpal seg-
ment 3; (36) wing; (37) spermathecae.
249
MATERIAL EXAMINED.
Java: Djakarta (see under S. perturbans). West Malaysia: Gunong Besout Forest Reserve, 1 2 (hypo-
pharynx and maxilla blade).
DIsTRIBUTION. China: Hainan (Leng, 1964 : 127); Mencheong (Yao & Wu, 1940 : 97); Aihsien, Paoting
(Yao & Wu, 19415: 76); Mang-shih (Yao & Wu, 1941c : 79). India: Saharanpur (Sinton, 1930b : 195);
Golaghat (Sinton, 1932c : 225); Sagar area (H. Trapido). Java: Jakarta (as above). Laos: Luang Prabang
(Parrot and Clastrier, 1952 : 153); Ventiane (Quate, 19626 : 256); Thahkek (Raynal, 19355: 241). Sri
Lanka: Kalgoda (Theodor, 1938 : 269). Thailand: Bangkok (Raynal & Gaschen, 1934a : 532). Vietnam
(North): Bac Muc, Coc Leu, Ha Giang, Lam, Nao Phu, Pho Moi, Phu Doan, Vinh Thuy, Yen Lay
(Raynal, 19355: 241); Hai Duong (Raynal, 1936 : 250); Cho Gan (Raynal & Gaschen, 1934a : 531).
Vietnam (South): 22 km south of Nah Trang (Quate, 19625 : 256); Kim Son (Raynal, 19355 : 241). West
Malaysia: Kuala Lumpur, Rantau Panjang light trap (Lewis, 1957 : 166); Batu (Quate, 1962a : 226).
Raynal (1936a : 350, 357) reported that P. stantoni seemed rare in India and was common in
Indo-China, being abundant between latitudes 20° and 30°.
UNGROUPED
Phlebotomus newsteadi Sinton
(Map 5)
Phlebotomus newsteadi Sinton, 1926 : 559; 1928a : 589 [2]; 1932a : 58; 1933e : 418; Theodor, 1948 : 418.
LECTOTYPE 3, InpIA: Kasauli; labelled ‘Type 3... Rabbit houses, Pasteur Institute, Kasauli,
Punjab, 28/8/25’ (BMNH), here designated [examined].
In the female the pharyngeal armature is well developed and the spermatheca is moniliform. In
the male there are two spines near the aedeagus, the paramere ends like a crochet-hook, three
of the spines on the style are apical or subapical, and the haltere is flattened.
Theodor placed this species provisionally in Euphlebotomus, but inspection of the lectotype
(below) suggests that it is outside the subgenera.
MATERIAL EXAMINED.
India: Kasauli, 1 ¢, as above.
DISTRIBUTION. India: Kasauli, Kurseong (Sinton, 1928c : 310).
P. newsteadi, like P. major, is a hill species (Sinton, 1932a).
Subgenus IDIOPHLEBOTOMUS Quate & Fairchild
Phlebotomus subgenus Idiophlebotomus Quate & Fairchild, 1961 :208; Theodor, 1965: 176; Lewis,
1973 : 162; Lewis & Lane, 1976 : 53; Abonnenc & Léger, 1976 : 76. Type-species : Phlebotomus asperulus
Quate & Fairchild, 1961, by original designation.
Idiophlebotomus Quate & Fairchild; Abonnenc, 1972 : 69, 75.
The cibarium has teeth, the pharynx is unarmed and the palp short. A pair of rods is associated
with the genital filaments. All the species are Oriental and most occur in caves. The redefinition
by Lewis & Lane should be altered as follows.
Pigment patch absent in all species but one. Antenna 3 with sub-basal swelling and beyond it a
slight depression bearing non-spatulate sensilla.
Phlebotomus (Idiophlebotomus) asperulus Quate & Fairchild
(Map 5)
Phlebotomus (Idiophlebotomus) asperulus Quate & Fairchild, 1961 : 208; Lewis & Lane, 1976 : 54. Holo-
type 3, WEesT MALAysIA (BPBM) [not examined].
The female has a cibarial median rod with large serrations, and the male has an apical spine on
the style which is markedly expanded.
250
DIsTRIBUTION. West Malaysia: Betis (Lewis & Wharton, 1963); Kuala Lumpur (Quate & Fairchild, 1961;
McClure et al., 1967 : 422).
Phlebotomus (Idiophlebotomus) erebicolus Quate
(Map 5)
Phlebotomus (Idiophlebotomus) erebicolus Quate, 1965:22; Lewis & Lane, 1976:57. Holotype 4,
PHILIPPINES (BPBM) [not examined].
The female has a cibarial median rod and a few granulose spicular teeth, and the male has three
spines on the style.
DISTRIBUTION. Philippines: Mainit Lake (Quate, 1965).
Phlebotomus (Idiophlebotomus) frondifer Lewis & Lane
(Map 5)
Phlebotomus (Idiophlebotomus) frondifer Lewis & Lane, 1976 : 57. Holotype 3, West MALAYSIA (BMNH)
[examined ].
The female has a median rod in the cibarium and very long parallel teeth, and the male has a
leaf-like aedeagus and four spines on the style.
DISTRIBUTION. West Malaysia: Gunong Besout Forest Reserve (Lewis & Lane, 1976).
Phlebotomus (Idiophlebotomus) pholetor Quate & Fairchild
(Map 5)
Phlebotomus (Idiophlebotomus) pholetor Quate & Fairchild, 1961 : 210; Lewis & Lane, 1976 : 57. Holotype
3, BORNEO: Sabah (BPBM) [not examined].
The female has a cibarial median rod and short teeth, and the male has five spines on the style.
DIsTRIBUTION. Borneo (Sabah): Gomantong (Quate & Fairchild, 1961). Philippines: Eran area (Quate,
1965 : 24).
Phlebotomus (Idiophlebotomus) sejunctus Quate
(Map 5)
Phlebotomus (Idiophlebotomus) sejunctus Quate, 1965: 20. Holotype 9°, PHiLippines (BPBM) [not ex-
amined ].
The female has a cibarium with no median rod and very long anterior teeth.
DIsTRIBUTION. Philippines: Mainit Lake (Quate, 1965).
Phlebotomus (Idiophlebotomus) stellae Quate
(Map 5)
Phlebotomus (Idiophlebotomus) stellae Quate, 1965:20; Lewis & Lane, 1976:59. Holotype 3,
PHILIPPINES (BPBM) [not examined].
The female has no median cibarial rod and very small teeth, and the male has five spines on
the style.
DIsTRIBUTION. Philippines: Minglanille (Quate, 1965).
2a
Phlebotomus (Idiophlebotomus) teshi Lewis sp. n.
(Map 5)
The female differs from other species of Jdiophlebotomus in having a cibarial pigment patch and
a median large short tooth.
?. Labrum 0-20 mm long, 0-08 length of wing, with two apical sensilla prominent, and reduced adoral
sensilla. Cibarium with chitinous arch far back, a very marked hind bulge, a large pigment patch, a
median very thick tooth and, on each side, about seven longitudinal rows of four to six teeth. Pharynx
with bulge twice as wide as fore part, and narrow hind opening with minute spicules. Hypopharynx
smooth with a few spicules at tip. Antenna 1 and 2 large, segment 3 = 0:58 mm long, 0-24 length of wing,
1-66 length of 4+ 5, 2-94 length of labrum, two strong ascoids on segments 3-16, that on 4 = 0-87 length
of segment and reaching next one, a papilla on 3-5. Mandible pointed, with small teeth and no submedian
pit. Maxilla toothless and shorter than mandible; palpal ratio 10: 14: 41 : 19: 20; with a rather diffuse
patch of slightly club-shaped sensilla. Thorax pale, without pleural hairs, inter-precoxal lobes broad.
Wing length 2-40 mm, 2:9 times width, R./R.i3 = 2°92, R; apex/R, = 0°54. Leg ratios: fore (1-04 mm),
10: 12:9; mid (0:96 mm), 10: 17:9; hind (1:00 mm), 10:19:14. Abdominal tergites 2-6 (as in P.
frondifer Lewis & Lane) with scattered erect hairs but few in mid line in hind halves of segments. Each
spermatheca having a short capsule with a very thick wall near the head, ducts thin-walled and indistinct;
furca with broad fore arm.
This species is named in appreciation of Dr R. B. Tesh’s studies of sandfly fever viruses and
Malaysian sandflies.
MATERIAL EXAMINED.
Holotype 2, Nepal: Pokhara, 18—27.ix.1965 (L. Quate), 910 m, Malaise trap, a little red blood in abdomen
(BPBM).
Phlebotomus (Idiophlebotomus) tubifer Lewis & Lane
(Map 5)
Phlebotomus (Idiophlebotomus) tubifer Lewis & Lane, 1976 : 59. Holotype 2, INDIA (BMNH) [examined].
The female has a median cibarial rod and teeth on radiating lines. The male has been discovered
by Dr G. B. Modi (1972, in letter).
DISTRIBUTION. India: Mahableshwar (Lewis & Lane, 1976).
Genus SERGENTOMYIA Franca & Parrot
Phlebotomus subgenus Newsteadia Franga, 1919 : 148. Type-species: Hebotomus minutus Rondani, 1843,
by subsequent designation of Franca, 1920 : 234. [Homonym of Newsteadia Green, 1902.]
Phlebotomus subgenus Sergentomyia Franca & Parrot, 1920 : 699; Fairchild, 1951 : 12; Quate, 1964 : 249.
[Replacement name for Newsteadia Franga. ]
[Neophlebotomus Franca & Parrot, 1920: 699. Incorrectly treated as synonym, now recognized as
subgenus. ]
Phlebotomus subgenus Prophlebotomus Franca & Parrot, 1921 : 281. Type-species: Hebotomus minutus
Rondani, 1843 : 265, by monotypy. [Synonymized by Theodor, 1948 : 88.]
Sergentomyia Franca & Parrot; Theodor, 1948 : 100; 1958 : 33; 1965 : 179; Perfil’ev, 1968 : 295.
Sergentomyia subgenus Sergentomyia Franca & Parrot; Theodor, 1948 : 101; 1958 : 36; Kirk & Lewis,
1951 : 409; Perfil’ev, 1968 : 214, 298; Abonnenc, 1972 : 69.
The cibarium has a definite row of teeth and a pigment patch. The style of the male has four
spines and an accessory seta. The genus is widely distributed in the Old World tropics and sub-
tropics.
In early papers on Indian sandflies, mainly before 1928, ‘P. minutus’ refers to one or more
species of the genus Sergentomyia, particularly the common S. babu. This was because the
taxonomic importance of cibarial teeth and spermathecae was unknown till 1926, and (Sinton,
1928b : 185) because S. babu was wrongly thought by Annandale (1911b) to be the Palaearctic
S. minuta (Parrot).
252
The musai-group
This group is here proposed for the single species S. musai.
Sergentomyia musai Lewis sp. n.
(Figs 38-50, Map 6)
The cibarium of the female has 12 long thread-like teeth and no fore teeth or pigment patch.
9. Labrum 0:22 mm long, 0-14 length of wing, with only two subapical sensilla and small adorals. Cibarium
with 12 long thread-like teeth, no pigment patch and a weak arch; salivary pump inapparent. Pharynx
damaged but showing lines and denticles. Hypopharynx with short row of nine teeth on each side.
Antenna 3 = 0:27 mm long, 0-17 length of wing, 1-01 length of 4+5, 1-20 length of labrum, two ascoids
on segments 3-15, that on 5 being 0-9 length of segment, papilla present on 5; hairs directed outward
and forward. Maxilla with no lateral teeth, 37 small but stout ventrals, and a dental depth of 0-05 mm;
palpal ratio 10 : 25 : 35 : 19 : 35; segment 3 with a few scattered spatulate sensilla, hair-like at some angles.
Scutum and pleuron pale, mesepisternum without hairs. Wing 1:58 mm long, 3:04 as long as wide,
R,/Ro+3 1°95, R, overlap/R, 0-66. Leg ratios: fore (0-62 mm), 10: 11-5 : —; mid (0-63 mm), 10 : 12:8 : 7:7;
hind (0-67 mm), 10: 14-4: -. Abdominal tergites 2-6 with many erect hairs and a few recumbent ones
near middle line. Spermatheca not fully visible but apparently simple.
ComMENT. This very unusual species differs from most Sergentomyia in its two labral subapical
sensilla, shape of cibarial teeth, hypopharynx, papilla on antenna 5, maxillary dentition, short
palpal segments 4 and 5 and nature of sensilla, and many abdominal erect hairs. S. musai
possesses several features of both Phlebotomus and Sergentomyia, and is placed, for the present,
among the ungrouped species of the latter. The hypopharynx and abdominal hairs of S. musai
may indicate relationship to the subgenus Parvidens Theodor & Mesghali. It is named after
Musa bin Long of the Arbovirus Unit, Kuala Lumpur, who collected many sandflies in West
Malaysia.
MATERIAL EXAMINED.
Holotype 2, Borneo: Sabah, Kalabakan area, 29.x.1962 (K. J. Kuncheria), forest camp 19 km north of
Kalabakan, 60 m (BPBM).
Subgenus SERGENTOMYIA Franca & Parrot
Sergentomyia subgenus Sergentomyia Franga & Parrot; Theodor, 1948: 101; 1958: 36; Perfil’ev,
1968 : 298.
The spermatheca is tubular with smooth sides and a wide duct, and the aedeagus is thick. The
subgenus is represented in the west of the Region.
Sergentomyia (Sergentomyia) dentata (Sinton)
(Figs 51, 52, Map 6)
Phlebotomus dentatus Sinton, 1933a : 869; 1933d : 228. Lectotype 2, PAKISTAN (BMNH), designated by
Lewis, 1967 : 25 [examined].
Sergentomyia (Sergentomyia) dentata (Sinton); Theodor, 1958 : 39 [in part]; Lewis, 1967 : 25; 1974 : 190.
In the cibarium of the female about four teeth on each side are much larger than the middle
ones.
2 (extra facts). Hypopharynx with distinct low rounded teeth. Mandible (as in S. punjabensis) with rather
blunt tip, and teeth large except near tip. Maxilla with ten broad lateral teeth, 20 distinct ventrals, and a
dental depth of 0:08 mm.
MATERIAL EXAMINED.
Pakistan: Landi Kotal.
233
emwrnw mw mg wm ew we ew ee’ ee ee ee ee ee eee Oe ee ee
a GS
Li
Sra)
=>
2)
Se
bh
foe)
+025
Figs 38-61 Sergentomyia species. 38-50, S. musai, 2: (38) labrocibarium; (39) tip of labrum;
(40) cibarium; (41) part of pharynx; (42, 43) tip of hypopharynx; (44) one side of antenna 6; (45)
maxilla; (46) palp segment 3; (47) wing; (48, 49) sockets of erect hairs on abdominal tergites 2-6
and on parts of 3 and 4; (50) tip of spermatheca. 51, 52, S. dentata, 2 (from Landi Kotal), labro-
cibarium and maxilla. 53-61, S. punjabensis, 2: (53) labrocibarium; (54-58) subapical sensillum
no. 2, adoral sensilla nos 1, 4 and 7, cibarial sensillum no. 5; (59) hypopharynx; (60) mandible;
(61) maxilla.
254
DIsTRIBUTION. Pakistan: Gwadi, Landi Kotal, Peshawar, Rawalpindi (Lewis, 1967 : 25); Quetta (Sinton,
1933a : 869).
Sergentomyia (Sergentomyia) punjabensis (Sinton)
(Figs 53-61, Map 6)
Phlebotomus antennatus Newstead; Sinton, 1927e : 31; 1933e : 421.
Phlebotomus minutus var. antennatus Newstead; Sinton, 1924a : 812; 1932a: 61 [9]; 1933e: 421; Qutu-
buddin, 1952 : 79.
Phlebotomus punjabensis Sinton, 1933e : 421. Conditional name available under ICZN Article 17(8).
No type-data (INDO-PAKISTAN, specially Punjab) [not examined].
Sergentomyia (Sergentomyia) punjabensis (Sinton); Theodor, 1948 : 109; Lewis, 1967 : 27; 1974b : 190.
Sergentomyia punjabensis (Sinton); Qutubuddin, 1951 : 36.
Phlebotomus antennatus var. deccanensis Qutubuddin, 1952 : 79. Syntype 3, INDIA (depository unknown)
[not examined]. [Synonymized by Lewis, 1967 : 27.]
The pharynx of the female is very wide posteriorly, and its armature has a deep acute hind notch
and many teeth of which the posterior ones are very small. The cibarial teeth are nearly equal
in size.
2 (extra facts). Labrum with adoral sensilla not differentiated into two groups, most of them about as
large as cibarials, some of each with long processes. Hypopharynx with distinct rounded main teeth, most
nearly twice as long as high. Mandible with large main teeth. Maxilla with strong teeth, five laterals and
22 ventrals, and a dental depth of 0-07 mm.
MATERIAL EXAMINED.
Pakistan: Shahzada (near Lahore).
DISTRIBUTION. India: Hardwar, Khandwa, Nagpur, Patiala (BMNH); Aurangabad (V. Dhanda);
Ajmer area, Jaipur, Jalor, Kota area, Sirohi area (Kaul et al., 1973 : 533); Jammu (Mitra & Roy,
1953a : 324); Hyderabad (Qutubuddin, 1944 : 208); Vellore (R. Reuben); Bikaner (Sharma et al., 1973c);
Bhavnagar, Bissamcuttack, Bombay, Calcutta, Chandigarh, Cochin area, Delhi, Itaunia, Junagadh,
Kamptee, Karnal, Madras, Parbitapur, Pinjaur, Pipariya, Pipli, Secunderabad, Singarama (Sinton’s
notes, as P. antennatus); Panchgani, Wai area (H. Trapido). Pakistan: Dehra Ismail Khan, Jhelum,
Khanki, Kohat area, Lahore, Peshawar, Rawalpindi, Said Pur (Lewis, 1967 : 28); Bannu, Lyallpur,
Sargodha, Tank (Sinton’s notes). Sri Lanka: Delft Island (Carter & Antonipulle, 1949 : 68).
Sinton (1932a) reported that ‘P. minutus var. antennatus’ (=S. punjabensis) had a more general
distribution over the Indo-Pakistan plains than did ‘P. minutus’ (=S. theodori).
Sergentomyia (Sergentomyia) theodori (Parrot)
[Phlebotomus minutus (Rondani, 1840 : 263); Adler & Theodor, 1926 : 403; Sinton, 1932a : 73; 1933e:
422; Parrot, 1942 : 330, 331. In part. ]
Phlebotomus (Prophlebotomus) theodori Parrot, 1942 : 332. [Replacement name for minutus auctorum
from Palestine. ]
Sergentomyia (Sergentomyia) theodori (Parrot); Theodor, 1948: 101; 1958:41; Theodor & Mesghali,
1964 : 294.
Sergentomyia (Sergentomyia) dentata (Sinton) form theodori; Lewis, 1974b : 191.
This belongs to a complex of closely related species and variants (Lewis, 19745), and its female
has a wider pharynx than in typical S. dentata.
Sergentomyia (Sergentomyia) theodori pashtunica Artemiev
(Map 6)
Phlebotomus minutus (Rondani); Sinton, 1932a : 61, 73 [in part]; 1933d: 421 [in part].
Sergentomyia (Sergentomyia) theodori (Parrot); Lewis, 1967 : 27.
ys
Sergentomyia (Sergentomyia) theodori pashtunica Artemiev, 1974b : 333. Syntypes 124 9, 209 ¢,
AFGHANISTAN (depository unknown) [not examined].
In the female, S. t. pashtunica has fewer cibarial teeth (22 = 18-24) and a shorter labrum
(0-14 = 0-13-0-16 mm) than the nominate form, and in the male differs by having a shorter
labrum (0-12-0-14) and shorter antenna 3 (0:10-0:11). Further study of variation and distribution
is needed to settle the status of S. theodori and its subspecies.
2 (extra facts). Fascicle much as in S. dentata, number of maxillary lateral teeth variable.
DISTRIBUTION. Pakistan: Dehra Ismail Khan, Kashmore, Landi Kotal, Larkana, Peshawar, Rawalpindi
(Lewis, 1967 : 27).
Sinton (1932a) stated that ‘P. minutus’ (= S. theodori) in the Indian subcontinent occurred chiefly
in the north-western plains.
Subgenus PARROTOMYIA Theodor
Sergentomyia africana group Theodor, 1948 : 101.
Sergentomyia subgenus Parrotomyia Theodor, 1958: 42; Perfil’ev, 1968 : 318; Artemiev, 1976 : 428
[some abdominal dorsal hairs erect in certain species]. Type-species: Phlebotomus africanus Newstead,
1912 : 363 (described as var.), by original designation.
A comb-like cibarial armature, lamp-glass-shaped pharynx, and elliptical capsular spermatheca
are characteristic of many species, but it is sometimes necessary to consider all the characters
given in couplet 69 of the key, and even they are not easily applicable in every case. The subgenus
is large and widely distributed. Some species have a short Rg.
The species are here placed in the africana, denticulata, babu and grekovi series. In the africana
series the ventral plate of the cibarium has no notch and the spermatheca is oblong. A few forms
(brevicaulis, himalayensis, mangana and queenslandi meridionalis) differ somewhat from the rest
in having a very high average, or sometimes maximum, number of cibarial hind teeth which may
also be very long with the outer ones directed slightly outward. Members of the denticulata-group,
which includes the Australasian S. spinosior (Quate & Quate), are like those of the africana-group
but have numerous parallel teeth so narrow (apparently less than a micron wide) as to be invisible
at most magnifications. The teeth are about as numerous as some in the africana-group which
are visible because they occupy a wider space and lean outward near the edges. In the babu-group
the ventral plate of the cibarium has a hind notch, and the spermatheca is oblong. In the grekovi-
group the spermatheca is nearly spherical.
Sergentomyia (Parrotomyia) sp. (A) Kaul, Dhanda & Modi
(Map 6)
Sergentomyia (Parrotomyia) sp. A Kaul, Dhanda & Modi, 1973 : 535 [3].
This species from the Kota district, India, may be a new one related to S. grekovi.
Sergentomyia (Parrotomyia) africana magna (Sinton) form asiatica Theodor stat. n.
(Map 6)
Phlebotomus africanus Newstead; Adler & Theodor, 1927 : 63 [Jericho]; Sinton, 1932a : 61; 1933e : 42.
Phlebotomus africanus var. asiaticus Theodor, 1933 : 541; 1952: 116 [relation to Israel form unknown].
Syntypes, IsRAEL (TC) [not examined].
Sergentomyia (Parrotomyia) africana var. asiatica (Theodor); Theodor, 1948 : 110.
Sergentomyia (Parrotomyia) africana asiatica (Theodor); Theodor, 1958 : 45; Lewis, 1967 : 28; Perfil’ev,
1968 : 331.
Phlebotomus africanus asiaticus Theodor; Abonnenc & Yvore, 1969 : 184; Bailly-Choumara, Abonnenc
& Pastre, 1971 : 454; Abonnenc, 1972 : 172.
Phlebotomus (Sergentomyia) africanus asiaticus Theodor; Abonnenc, 1972 : 172.
256
The cibarium of the female has 45-50 teeth in a concave row and no central patch of fore teeth,
and the pigment patch is concave posteriorly and has a simple forward process, and there is no
notch in the ventral plate. The pharynx has long teeth and the spermatheca is oblong.
The S. africana complex is mainly African and includes a number of forms which have a
comb-like cibarial armature and oblong spermatheca, and have been variously treated as several
species or as one by Quate (1964). His citation of 36 name-combinations occupies 137 lines and
shows the confusion which existed for many years. He, Lewis (19745) and others have empha-
sized the degree of intergradation between some named forms. There is so much variation that
anyone with a large collection at his disposal could, if so minded, create still more names for
local variants. For these reasons I regard S. a. magna as no more than a subspecies.
No certain specimens of the Pakistan form are available, but it probably belongs to a small
northern form found in Morocco, Israel and India, which Abonnenc & Yvore (1969 : 184, 185)
and Abonnenc (1972 : 172) treated as a subspecies of africanus. Rioux et al. (1975) regarded it
as infra-subspecific. For the present I am treating the Pakistan form as a form of S. a. magna.
MATERIAL EXAMINED.
India: Palod (1934, M. O. T. Iyengar), 1 2 with about 38 hind teeth and a very concave pigment patch
may be related.
DISTRIBUTION. Pakistan: Kandhkot, Rhedia, Shikarpur (Lewis, 1967 : 28).
Sergentomyia (Parrotomyia) sp. (B) Kaul, Dhanda & Modi
(Map 6)
Sergentomyia (Parrotomyia) sp. B Kaul, Dhanda & Modi, 1973 : 536 [3].
Males from Kota district, India, may be a new species. It is provisionally put in the africana-
group.
Sergentomyia (Parrotomyia) babu (Annandale)
Phlebotomus babu Annandale, 19106 : 49.
The cibarium of the female has a deep notch in the ventral wall and 24-50 teeth.
Sinton (1932a : 60; 1933e : 422) pointed out the close affinity between S. babu, S. baghdadis
and S. shorttii but retained their specific status in view of their morphology and distribution.
S. babu and S. baghdadis overlap in a considerable area with little intergradation, but many
intermediate females, classed here as a S. babu variant, have been found in the Sagar (Shimoga)
area. Females from Nander and Parbhani, intermediate between S. baghdadis and S. shorttii,
are classed here as a S. baghdadis variant; typical S. baghdadis may not occur there. S. babu and
S. baghdadis, and now S. shorttii, are known to overlap in large enough areas to be regarded as
species, but their true status must await further knowledge.
Sergentomyia (Parrotomyia) babu babu (Annandale)
(Map 6)
Phlebotomus sp., Howlett in Maxwell-Lefroy, 1909 : 559 [?].
Phlebotomus babu Annandale, 1910b: 49; Sinton, 1927e:31; 19285: 185; 1928c: 314; 1932a: 60;
1933d : 422; Theodor, 1938 : 264 [24-34 cibarial teeth in 2]. Lectotype 3, INDIA (Zoological Survey of
India, Calcutta), designated by Quate, 1962c : 158 [not examined].
Phlebotomus babu var. niger Annandale, 1911c : 320; Sinton, 1928c : 315; 1932a: 60. Type, INDIA (be-
lieved by Quate to be lost) [not examined]. [Synonymized by Sinton (1932a) and Quate (1962c). ]
Phliebotomus minutus var. niger Annandale; Sinton, 1924a : 812; 1927d : 25; 1927e : 31.
Phlebotomus (Prophlebotomus) babu Annandale; Parrot, 1940 : 311.
Phlebotomus thapari Mitra & Roy, 1952 : 88. Holotype 2, INpIA (depository unknown) [not examined].
Syn. n.
Zot
Phlebotomus (Sergentomyia) babu var. niger Annandale; Quate, 1962c : 158.
Phlebotomus (Sergentomyia) babu Annandale; Quate, 1962c : 157.
Sergentomyia (Parrotomyia) babu (Annandale); Lewis, 1967 : 88; Artemiev, 1976 : 422.
The cibarium of the female has 24-34 teeth. In an aberrant female from India in the BMNH,
marked K1 by Sinton, vein R, is absent in one wing and very short in the other.
The form thapari, based on a single female, appears to be a synonym, and some of its features
due to artefacts and interpretation.
DISTRIBUTION. India: Abu Mount, Badam Pahor, Delhi, Krishnagar, Nagpur, Osmanabad, Pinjaur,
Saharanpur (BMNH); Aurangabad, Jalna, Patan (Farooq & Qutubuddin, 1945 : 85); Ajmer, Sambhar
(Jaswant Singh, 1933); Tibi area (N. L. Kalra); Jalor, Kota, Sirohi (Kaul et al., 1973 : 532); Calcutta,
Poona, Pusa (Lewis, 1967 : 30); Allahabad, Asansol, Igatpuri, Palod Port Canning, Purnea, Rajmahal,
Rambha, Trivandrum (Quate, 1962c: 158); Bikaner (Sharma et al., 1973c); Bombay, Chindwara,
Faizabad, Itarsi, Itaunia, Junaghad, Kamptee, Karnal, Kurseong, Mhow, Nabinagar, Panchmarhi,
Patna (Orissa), Pipariya, Rajkot, Singanama, Titilagarh (Sinton’s notes); Sagar (Shimoga) area (some
typical, many with about 22 cibarial teeth), Wai (H. Trapido); Naini Tal (Wattal et a/., 1967). Pakistan:
Cherat, Gilgit, Lahore, Kotal, Rawalpindi, Taxla (Lewis, 1967 : 30); Khanki (Sinton’s notes).
According to Sinton (1928c : 315; 1932a), S. babu was the main component of ‘P. minutus’ till
1928, and was evidently widely distributed over the Indo-Pakistan plains and foothills. Many
localities of this common species are omitted from the above list.
Sergentomyia (Parrotomyia) babu insularis (Theodor)
(Map 6)
Phlebotomus babu var. insularis Theodor, 1938 : 264. Syntypes 6 2, 7 3, SR1 LANKA (TC) [not examined].
Sergentomyia (Sergentomyia) babu var. insularis Theodor; Theodor, 1948 : 110.
Phlebotomus babu Annandale or ally [?]; Carter & Antonipulle, 1949 : 68.
Sergentomyia (Sergentomyia) babu insularis Theodor; Lewis, 1973a : 249.
The cibarial teeth of the female number 45-50. There is much variation in the number of teeth
in some species of Parrotomyia, and this form is treated as a subspecies partly owing to its
geographic isolation.
DISTRIBUTION. Sri Lanka: Depanama, Maharamgama, Pannipitiya (Theodor, 1938 : 266).
Sergentomyia (Parrotomyia) baghdadis (Adler & Theodor)
(Map 7)
Phlebotomus baghdadis Adler & Theodor, 1929: 281; Sinton, 1932a: 60; 1933e:422. 23 syntypes,
IRAQ (TC) [not examined].
Sergentomyia (Parrotomyia) baghdadis (Adler & Theodor); Lewis, 1967 : 30; Artemiev, 19765 : 424.
The female has about 16-20 cibarial teeth, a notch near them, and a weak pharyngeal armature.
DIsTRIBUTION. India: Nander and Parbhani (with 16-18 rather uniform cibarial teeth and a small or very
small notch, V. Dhanda); Riasi (Jacob & Kalra, 1951 : 325); Ajmer, Sambhar (Jaswant Singh, 1933);
Jaipur, Kota (Kaul et al., 1973 : 532); Punch, Rajouri (Mitra, 1959 : 62); Ferozepore, Karnal (Sinton’s
notes). Pakistan: Sukkur (BMNH); Bannu, Dehra Ismail Khan, Gujranwala, Gujrat, Jandola, Jhelum
Kandhkot, Kashmore, Lahore, Landi Kotal, Larkana, Lyallpur, Mir Muhammad, Pano Agil, Peshawar,
Rawalpindi, Rohri, Said Pur, Sarghoda, Shikarpur, Tank, Taxla (Lewis, 1967 : 31).
In the Indian subcontinent this species occurs in the west and north-west (Sinton, 1932a).
258
Sergentomyia (Parrotomyia) barraudi (Sinton)
(Figs 62-69, Map 7)
Phlebotomus barraudi Sinton, 1929a: 716; 1932a:61; 1933e:422; Raynal & Gaschen, 19345: 559;
1935a : 113; Raynal, 19355 : 285; Theodor, 1933 : 542 [variety of africana ?]; 1938 : 268 [variation];
Tonnoir, 1935 : 142; Yao & Wu, 19416 : 78. Syntypes 2 3, INDIA (depository unknown) [not examined].
Phlebotomus (Prophlebotomus) barraudi Sinton; Parrot, 1937: 120; 1940:311; 1946: 70; Parrot &
Clastrier, 1952 : 158.
Phlebotomus barraudi var. siamensis Causey, 1938 : 488. Syntypes 2 9, THAILAND (depository unknown)
[not examined]. [Synonymized by Quate, 19625: 261.]
Phlebotomus barraudi var. kwangsiensis Yao & Wu, 1941a:67; 19415: 77. Syntypes 2 92, 1 g, CHINA
(depository unknown) [not examined]. [Synonymized by Quate, 19625 : 261.]
Phlebotomus barraudi var. siulamensis Chen & Hsu, 1955 : 302; Leng, 1964 : 124, 127. Syntypes, CHINA:
Samshuihsien (Hua Nan Medical College, Canton) [not examined]. Syn. n.
Sergentomyia (Parrotomyia) barraudi (Sinton); Theodor, 1958 : 44; Lewis & Dyce, 1976 : 208.
Phlebotomus (Sergentomyia) barraudi Sinton; Quate, 1962b : 261; Cates & Lien, 1970 : 535.
The normally bifid shape of the pigment patch of the female is a characteristic feature of this
species. The number of cibarial teeth in the female is recorded by Causey, Chen & Hsu, Lewis &
Dyce, Quate, Raynal (1935), Sinton (1929) and Theodor, and ranges from about 40 to about
70, being particularly high in some eastern areas. The following note is based on females from
West Malaysia.
9. Tip of labrum small and median apical sensilla prominent. Hypopharyngeal teeth delicate. Mandible
pointed. Maxilla with 2-8 (2-4) lateral nodular teeth and 35-4 (35-36) ventrals (n = 5); dental depth
0-10 mm; lateral teeth distinctly smaller in West Malaysia than in India and East Timor.
Professor O. Theodor informed me (1971, in letter) that specimens from Taiwan resemble the
type form in general but differ in having a shorter antenna 3 in the female and a short wide
pigment patch like that of some Chinese specimens. He remarked that the species seemed to
have some local forms and that more material was necessary for assessing their status.
MATERIAL EXAMINED.
Hong Kong: Hong Kong (G. W. Chau), 1 2 (with 64 hind teeth and faint or no pigment patch, and much
indented cibarium and pharynx), 2 2 (with about 70 teeth). Java: Semarang, ii.1910 (E. Jacobson), 1 9
(with some 90 cibarial teeth about 6 um wide, a partly divided process of the pigment patch, and a
nodular hind margin of the cibarial ventral plate; it may be related to S. barraudi). West Malaysia:
Gunong Besout area.
DISTRIBUTION. Bangladesh: Dhurmakura (Sinton’s notes). Burma: Mezali (Sinton’s notes). Cambodia:
Phnom Penh (Parrot & Clastrier, 1952 : 153). China: Sainan, Suilam (Chen & Hsu, 1955 : 302); Hainan
(Leng, 1964: 127); Se-Tchouen (Gaschen, 1934: 890; Raynal, 19355:290); Mang-shih, Nanning,
Tienpao (Yao & Wu, 1941a:67; 1941c:79). Hong Kong: (as above). India: Golaghat (Sinton,
1929a : 716); Gauhati (Sinton’s notes). Laos: Vientiane (Quate, 19625 : 261); Xieng Khouang (Raynal,
19355 : 290). Nusa Tenggara: Dili, Sumbawa (Lewis & Dyce, 1976 : 208). Japan: Ryukyu Retto, Iriomote
Island (Ohara), 1963 (G. A. Samuelson), 1 2; R. R., Ishigaki Island (Banna), 1963 and 1964 (J. L. Gressitt
& G. A. Samuelson, U.S.-Japan Cooperative Scientific Programme), via BPBM, 18 9, 1 3, the only species
found. Taiwan: Hengchun (C. M. Yoshimoto); Hualien area, Tapei area, Taitung (Cates & Lien, 1970:
537, T. C. Maa); Kuraru (C. M. Yoshimoto). Thailand: Bangkok (Causey, 1938 : 388); Phra Phuttabat
area (Quate, 19625 : 261). Vietnam (North): Cho Ganh (Raynal, 19355 : 290). Vietnam (South): Di Linh
(Quate, 19625 : 261); Duc Pho (Raynal, 19355 : 290). West Malaysia: Gunong Besout Forest Reserve
(A. B. Knudsen & colleagues); Klang, Ulu Gombak (D. J. Lewis); Kuantan, Tanjong Rabok (A. Rudnick).
In Indo-Burma Sinton (1932a) knew S. barraudi only from Assam and Burma. In Indo-China
Raynal (1936a : 366) found it between 15° and 25° north, where it was abundant at 1200 and
1600 m in particular, and he regarded it as a hill species.
Sergentomyia (Parrotomyia) bigtii (Manalang)
(Map 7)
Phlebotomus bigtii Manalang, 1931 : 356; Raynal, 1935a : photograph 18; Quate & Rosario, 1962 : 797;
259
Quate, 1965 : 28. Lectotype 9, PHILIPPINES (on loan to BPBM), designated by Quate, 1965 : 28 [not
examined ].
The female of this large species has 10-12 bent pointed teeth, a rather narrow dark pigment
patch and a thick mass of pharyngeal teeth.
DISTRIBUTION. Philippines: Bigti, Malinao (Quate, 1965 : 28).
Sergentomyia (Parrotomyia) brevicaulis (Quate)
(Map 7)
Phlebotomus (Sergentomyia) brevicaulis Quate, 1962b: 256. Holotype 9°, VIETNAM (SOUTH) (BPBM)
[not examined].
This is a large species with, in the female, about 73 cibarial hind teeth (number taken from
figure), about 50 fore teeth in one row, a large pigment patch and long antenna 3 and Rg.
DIsTRIBUTION. Vietnam (South): Di Linh area (Quate, 19626 : 259).
Sergentomyia (Parrotomyia) brevinervis (Quate & Fairchild)
(Map 7)
Phlebotomus (Sergentomyia) brevinervis Quate & Fairchild, 1961 : 214 [¢]. Holotype 3, BoRNEo (BPBM)
[not examined ].
In the male there are 14 cibarial hind teeth and 13 fore teeth, no pigment patch, short antenna 3
and a narrow wing with short R,. The authors compared the male with S. babu. The species is
placed provisionally in the africana-group.
DISTRIBUTION. Borneo (Sabah): Ranau (Quate & Fairchild, 1961 : 214).
Sergentomyia (Parrotomyia) bukidnonis (Quate)
(Map 7)
Phlebotomus (Sergentomyia) bukidnonis Quate, 1965 : 33. Holotype 2, PHILIPPINES (BPBM) [not examined].
In the female of this small species the broad cibarium has 30-34 sharp teeth in a crown-like row
and 18-24 fore teeth in one row, the pharynx is unarmed, antenna 3 is short, palpal segment 4
is long, and R, is longer than Rg, 5.
DISTRIBUTION. Philippines: Cuernos de Negros, Katanglad Mountain (Quate, 1965 : 33).
Sergentomyia (Parrotomyia) dayapensis (Manalang)
(Map 7)
Phlebotomus dayapensis Manalang, 1931 : 358; Raynal, 1935a: photograph 17. Holotype 9, PHILIPPINES
(destroyed, according to Quate & Rosario) [not examined].
Phlebotomus (Sergentomyia) dayapensis Manalang; Quate & Rosario, 1962 : 793 [specificity of ¢ ques-
tioned ].
In the female the cibarium has some 24 teeth and a broad pigment patch, the pharynx is unarmed,
and R, equals Ry,s.
DISTRIBUTION. Philippines: Dayap (Manalang, 1931 : 355).
260
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Figs 62-79 Sergentomyia species. 62-64, S. barraudi, 2 (from Gunong Besout area), labral tip,
maxilla and spermatheca. 65-69, S. kauli, 2: (65, 66) cibarium; (67, 68) pharynx; (69) sperma-
theca. 70-73, S. modii, 2: (70) cibarium; (71, 72) pharynx; (73) spermatheca. 74-79, S. rudnicki, ?:
(74-76) cibarium, hind teeth, and fore teeth after KOH treatment; (77, 78) pharynx; (79) sperma-
theca.
261
Sergentomyia (Parrotomyia) denticulata (Quate & Fairchild)
(Map 7)
Phlebotomus (Sergentomyia) denticulatus Quate & Fairchild, 1961 : 216. Holotype 9, BoRNEo (BPBM)
[not examined ].
The cibarium of the female has about 90 teeth pointing upward, each one being very narrow
(apparently less than 1 ~m wide) so that at most magnifications the armature looks like a sheet
of chitin. The fore teeth number about 20, the pharynx is unarmed, the scutum is brown, and
R,/Ry,3 is about 1-9. Labrum 0-09 length of wing, in a female paratype.
A female from Sepilok is provisionally placed in this species although its labrum is 1-25 length
of wing and the cibarial teeth are scarcely visible, perhaps obscured by food or other structures.
DIsTRIBUTION. Borneo (Sabah): Sepilok (D. J. Lewis, 21 2); Gomantong Caves, Ranau (Quate & Fair-
child, 1961 : 217).
Sergentomyia (Parrotomyia) franciscana (Quate)
(Map 7)
Phlebotomus (Sergentomyia) franciscanus Quate, 1965 : 33. Holotype 9°, PHILIPPINES (BPBM) [not ex-
amined ].
In the female the cibarium has 24—26 teeth in a crown-like row and 14-18 fore teeth in one row,
the pharynx is unarmed, and R,j is shorter than Rg, <3.
DIsTRIBUTION. Philippines: San Francisco area (Quate, 1965 : 33).
Sergentomyia (Parrotomyia) grekovi (Khodukin)
(Map 7)
Phlebotomus grekovi Khodukin, 1929: 101. Syntypes 9, 3, U.S.S.R. (depository unknown) [not ex-
amined ].
Sergentomyia (Parrotomyia) grekovi (Khodukin); Theodor, 1958:44; Lewis, 1967:32; Perfil’ev,
1968 : 319; Artemiev, 1976b : 425.
In the female the cibarium has 18-35 teeth, possibly indicating more than one form, and the
spermatheca is nearly spherical. Artemiev, in Afghanistan, noted much variation in numbers of
teeth, and the presence of some erect hairs on certain abdominal tergites.
DISTRIBUTION. Pakistan: Gilgit (Lewis, 1967 : 34).
Sergentomyia (Parrotomyia) himalayensis (Annandale)
(Map 7)
Phlebotomus himalayensis Annandale, 1910b:50; Sinton, 1924a:811; 1924b:321; 19315: 828;
1932a: 62 [2]; 1933e: 421. Lectotype 9, INDIA (Zoological Survey of India), designated by Quate,
1962c : 158 [not examined].
Phlebotomus (Sergentomyia) himalayensis Annandale; Quate, 1962c : 158.
The female has about 80-100 cibarial hind teeth in a slightly concave line and a triradiate pig-
ment patch, and many short stout pharyngeal teeth. The male has about 40 cibarial teeth and its
style bears at or near the tip three large spines, one small spine and the usual seta.
MATERIAL EXAMINED.
Three females from the Jog-Sagar area of India (H. Trapido), with 66 hind teeth and a roundly truncated
process on the pigment patch, may be this species or a related form.
262
DIsTRIBUTION. India: Bhowali, Naini Tal (Brunetti, 1912 : 205; Sinton, 1928c : 321); Kurseong (1520 m
in eastern Himalayas, Sinton, 19245 : 817).
Sergentomyia (Parrotomyia) kauli Lewis sp. n.
(Map 7)
The female slightly resembles S. africana, but has a narrow pigment patch, and very short upper
pharyngeal teeth and long lower ones.
9. Labrum 0-23 (0:22-0:25) mm long, 0-11 (0:10-0:12) length of wing. Cibarium with 26-32 teeth, fore
teeth if present hidden by pigment patch; patch dark with broad anterior process. Pharynx with long
teeth. Antenna 3 = 0-23 (0:22-0:24) mm long, 0-12 (0-11-0-12) length of wing, 1-08 (1-03-1-14) length
of 4+5, 1-01 (0:90-1-09) length of labrum, ascoid on 4 about 0-43 length of segment and not reaching
next one. Wing length 1-95 (1:84-2:16) mm, 3-5 times width, R,/R2+i3 1°82 (1:66-2:11), R, tip/R, 0-65
(0:58-0:68). Spermatheca thick-walled and oblong with straightening very near tip.
ComMENTS. The specimens from Saharanpur had been provisionally labelled ‘P. malabaricus’ and
presented to the BMNH by J. A. Sinton. Therefore they were first presumed to be S. modii
which had been regarded as the female of S. malabarica till the true female was found. S. kauli
is shown to be a separate species, however, by the rounded hind lateral walls of the cibarium,
more numerous hind teeth, lack of conspicuous irregular fore teeth, broad forward process of
the pigment patch, shorter antenna 3 and ascoid, and relatively short Rg.
MATERIAL EXAMINED.
Holotype 9, India: Ganjam, 21.xi.1972 (H. N. Kaul) (BMNH).
Paratypes. India: Ramanagar, 28.viii.1970 (H. R. Bhat), 400m, 1 9; Saharanpur, ix-x.1927 (J. A.
Sinton), 4 2. (All in BMNH.)
Non-paratypic material. India: Munikireti, 6.iii.1973 (H. R. Bhat), 1 9 (BMNH).
Sergentomyia (Parrotomyia) mangana (Manalang)
(Map 7)
Phlebotomus mangana Manalang, 1930c : 283; Tonnoir, 1935 : 142. Lectotype 9, PHILIPPINES (BPBM),
designated by Quate & Rosario [not examined].
Phlebotomus (Sergentomyia) mangana Manalang; Quate & Rosario, 1962 : 793.
The cibarium of the female has 65-70 hind teeth, two rows of 20-25 fore teeth, and between
them six to eight larger teeth; palpal segment | is long.
DIsTRIBUTION. Philippines: Tunkong Manga (Manalang, 1930c : 283); Bigti, La Mesa dam area, Los
Bajfios, San José del Monte, San Mateo, Silang, Tala (Quate & Rosario, 1962 : 794).
Sergentomyia (Parrotomyia) modii Lewis sp. n.
(Figs 70-73, Map 7)
Phlebotomus sp. Sinton, 1927a : 933 [measurements].
[Phlebotomus malabaricus Annandale; Sinton, 1927d:25 [spermatheca]; 1928c : 321; 1932a:61, 74
[cibarium and pharynx]. Misidentification. ]
_ S. modii is distinguished from S. kauli by having fewer hind teeth, and from S. yoshimotoi by
having a pigment patch. Aspects of S. modii are noted under S. malabarica
9. Labrum 0-25 (0:24-0:26) mm long, 0-12 (0-11-0-12) length of wing. Cibarium with 17 hind teeth and a
few irregular fore teeth; pigment patch with long, rather narrow, forward extension. Pharynx with long
teeth. Hypopharynx smooth. Antenna 3 = 0°31 (0:30-0:32) mm long, 0-14 (0:14-0:15) length of 4+5,
1-25 (1:22-1:27) length of labrum; ascoid on 4 = 0-74-0-82 length of segment and extending to 5. Palpal
ratio 10:22: 32:35:72. Wing length 2:16 (2:16-2:16) mm, 3-3 times width, R,/R:3 2°97 (2:48-3:45),
R, apex/R, 0-79 (0:75-0-82). Spermatheca oblong with well-defined smooth wall.
263
MATERIAL EXAMINED.
Holotype 2, India: Gologhat, v.1925 (P. J. Barraud) (BMNH).
Paratype. 1 2, same data as holotype but 22.v.1925 (BMNH).
Non-paratypic material. A female with 28 hind teeth from Banhasa, India, may be related to this
species.
Sergentomyia (Parrotomyia) palestinensis (Adler & Theodor)
(Map 8)
Phlebotomus palestinensis Adler & Theodor, 1927 : 64. Holotype 9, IsRAEL (TC) [not examined].
Sergentomyia (Parrotomyia) palestinensis (Adler & Theodor); Theodor, 1958 : 44; Lewis, 1967 : 34;
Perfil’ev, 1968 : 326; Artemiev, 1976b : 427.
The female has 15-22 cibarial teeth in a straight line and a nearly spherical spermatheca. Artemiev
noted the close relationship of this species to S. sogdiana (Parrot).
DIsTRIBUTION. Pakistan: Peshawar (Lewis, 1967 : 34).
Sergentomyia (Parrotomyia) queenslandi (Hill)
Phlebotomus queenslandi Hill, 1923 : 83. Syntypes 4 9, 4 3, AUSTRALIA (National Museum, Melbourne)
[not examined ].
The pharynx of the female has hair-like scales and about 45-80 cibarial teeth.
Sergentomyia (Parrotomyia) queenslandi meridionalis (Tonnoir)
(Map 8)
Phlebotomus queenslandi meridionalis Tonnoir, 1935: 140. Syntypes 8 92, 3, AUSTRALIA (Division of
Economic Entomology, Melbourne) [not examined].
Sergentomyia (Sergentomyia) queenslandi var. meridionalis (Tonnoir); Theodor, 1948 : 111.
Sergentomyia (Parrotomyia) queenslandi meridionalis (Tonnoir); Lewis & Dyce, 1976 : 210.
The female has about 80 cibarial hind teeth. This form is treated as a subspecies till more is known
of its variation and distribution.
DISTRIBUTION. Nusa Tenggara: Tafara Cape (East Timor, Lewis & Dyce, 1976 : 210).
Sergentomyia (Parrotomyia) rudnicki Lewis sp. n.
(Figs 74-89, Map 8)
In the female of this pale species the cibarial teeth are so narrow as to be scarcely visible, the
pigment patch is broad and dark, and the pharyngeal teeth are strong.
The female differs from S$. mangana in having a short labrum, a shorter row of cibarial teeth
(0-056 mm in mangana), narrower teeth (0-8 um in mangana), two rows of fore teeth, and simple
outline to the forward process of the pigment patch.
The female of S. rudnicki differs from S. queenslandi meridionalis in having a shorter tooth-row
(0-072 mm in S. q. meridionalis) and narrower teeth.
The female of S. rudnicki differs from the Australasian (West Irian) S. spinosior (Quate &
Quate) in having a wider cibarial armature, shorter pharyngeal ventral teeth, and a long antenna
3 which is distinctly longer than 4+5.
2. Labrum 0-16 (0:15-0:18) mm long, 0-10 (0:09-0:10) length of wing. Cibarium with a row, about 0°43
mm long, of some 90 scarcely visible teeth, each about 0-05 um wide; two even rows of about 22 fore
teeth (seen in macerated specimens) hidden by dark brown posteriorly convex pigment patch; forward
process of patch with irregular margin and merging into suture. Pharynx with thick walls, and straight
264
sides beyond main bulge, and long narrow teeth, the dorsal ones appearing less long because they are seen
foreshortened or end-on. Hypopharynx with no teeth or undulations. Antenna 3 = 0-29 (0:26-0:33) mm
long, 0-17 (0:16-0:19) length of wing, 0-12 (0:12-0:13) length of 4+.5, 1-80 (1-66—1-92) length of labrum,
two ascoids on segments 3-15, that on 4 = 0-395 length of segment, no papilla on 5. Maxilla with nine
lateral teeth, 19 ventrals and a dental depth of 0-06 mm;; palpal ratio 10 : 21 : 38 : 40 : 80; clavate sensilla
not very numerous, on basal half of 3. Scutum, pleuron and whole body pale. Wing length 1-69 (1 -63—1-74)
mm, 3-3 times width, R,/R..3 1-61 (1:41-2:00), R, overlap/R, 0-59 (0:47-0:71). Abdominal tergites 4-6
with about one, two and four erect hairs respectively. Spermatheca oblong and sometimes slightly
wrinkled, with broad knob and shallow pit, and very thin ducts.
3. Labrum 0-15 (0:14-0:16) mm long, 0-10 (0-:09-0-10) length of wing. Cibarium with about 26 hind
teeth and a rather dark, or pale, subtriangular pigment patch having a thin forward process. Pharynx
rather as in 2 but less armed. Antenna 3 = 0-32 (0:29-0:38) mm long, 0-21 (0:20-0:24) length of wing,
1-16 (1:07-1:29) length of 4+5, 2-18 (2:04-2:40) length of labrum, one ascoid on 3-15. Wing length
1-52 (1-45-1-60) mm, 3-7 times width, R,/R.2,3 1:25 (0-89-1-50), R, overlap/R, 0-57 (0-46-0-67). Aedeagus
narrow and tapering to a blunt tip. Filament about 3-5 length of pump. Paramere beaked. Style with two
spines terminal and two subterminal, and a seta proximal to them.
MATERIAL EXAMINED.
Holotype 2, West Malaysia: Ulu Gombak, 30.x.1968 (Abu Hassan bin Omar) (BMNH). Ten of each sex
measured.
Paratypes. West Malaysia: as for holotype, 26.viii-30.ix.1968, 14 9, 3 ¢; 29.ii.1972 (D. J. Lewis), 1 9,
8 3; Ulu Langat Forest Reserve, 25.vii.1968 (A. Rudnick), 1 9, 1 3. (Allin BMNH.)
Sergentomyia (Parrotomyia) shorttii (Adler & Theodor)
(Map 8)
Phlebotomus shorttii Adler & Theodor, 1927 : 65 [palp formula variable]; Sinton, 1928c : 317; 1932a : 60;
1933e : 422. Syntypes 8 9, 8 3, INDIA (TC) [not examined].
Phlebotomus (Prophlebotomus) shorttii Adler & Theodor; Parrot, 1937 : 120.
Sergentomyia (Parrotomyia) shorttii (Adler & Theodor); Lewis, 1967 : 31; Perfil’ev, 1968 : 13.
The female is distinguished by having a notch near the cibarial teeth which number 10-14.
DISTRIBUTION. Bangladesh: Dacca (BMNH); Bongong, Kaoraid, Sylhet (Sinton’s notes). Burma:
Rangoon (Sinton, 1927e : 31). India: Golaghat (Adler & Theodor, 1927 : 66); Gauhati, Sukna (BMNH);
Kota (Kaul ef al., 1972 : 533); Soraipani (1938, R. C. Muirhead-Thomson); Hyderabad (Qutubuddin,
1944 : 208); Naini Tal area (Rao et al., 1973); Calcutta, Jog-Sagar area, Kannur, Konehosur, Kumsi
(H. Trapido). Pakistan: Taxla (Lewis, 1967); Lahore (George, 1970).
Sinton (1928c) found that S. shorttii represented the S. babu complex in Assam, and (1932a)
knew the former from the north-eastern parts of India and from Burma.
Sergentomyia (Parrotomyia) spinifaucis (Quate)
(Map 8)
Phlebotomus (Sergentomyia) spinifaucis Quate, 1965 : 28. Holotype 2, PHILIPPINES (BPBM) [not examined].
In the female the cibarium has 50-60 teeth in a row concave medially and straight or convex
laterally, and 10-12 fore teeth of which the median six are very large; antenna 3 is short and
R,/Ro,3 is about two. S. mangana is somewhat similar but is larger, has a differently shaped
_ tooth-row and a longer antenna 3.
DisTRIBUTION. Philippines: Tarumpitao (Quate, 1965 : 28).
Sergentomyia (Parrotomyia) timorica Lewis & Dyce
(Map 8)
Sergentomyia (Parrotomyia) timorica Lewis & Dyce, 1976 : 210. Holotype 2, Timor (Australian National
Insect Collection) [examined ].
265
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ee 10/25,
SRwet ee
a ee “
ey eee > —_—_—_—_—_——
tepeg O 2, ‘ s
0? tapy get
Figs 80-92 Sergentomyia species. 80, S. rudnicki, 3, terminalia. 81-89, S. torrechantei, 2: (81) labro-
cibarium; (82) cibarium; (83, 84) pharynx; (85, 86) maxilla and palp; (87) wing; (88) inter-precoxal
lobe; (89) spermatheca. 90-92, S. balica (from Carey Island): (90, 91) 9, cibarium and pharynx;
(92) 3, terminalia.
266
In the female the cibarium has about 60 (50-65) teeth and a slender pharynx with few spicules,
and R,/R»,,5 is 0.70 (0:24-0:92).
DiIsTRIBUTION. Nusa Tenggara: Dili, Kupang area, Pedang Bay (?), Tafara Cape (Lewis & Dyce,
1976 : 210).
Sergentomyia (Parrotomyia) torrechantei (Manalang)
(Map 8)
Phlebotomus torrechantei Manalang, 1931 : 361. Syntypes 2 9, 1 3, PHILIPPINES (destroyed) [not examined].
Phlebotomus (Sergentomyia) torrechantei Manalang; Quate & Rosario, 1962 : 797.
The original small figures of the female show about 30 cibarial hind teeth, dark walls, a marked
pigment patch and long pharyngeal teeth. Quate & Rosario comment that this species appears
close to S. mangana which has numerous fine cibarial teeth, an armed pharynx and similar wing
venation, but that S. torrechantei has a shorter palpal segment 4 and a differently shaped pigment
patch. The following description is from females believed to be this species.
9. Eye 0-58 length of head. Labrum 0-17 (0-:17-0-18) mm long, 0-09 (0-09-0-09) length of wing, with
well-marked sensilla of which the distal ones are broad. Cibarium with about 48 teeth in a slightly convex
row, dark, rather narrow, pigment patch with strong forward process, a brown band on ventral wall
partly hiding patch, dark sides and a scarcely visible chitinous arch. Pharynx with strong armature of
pointed teeth, those on ventral plates conspicuous. Hypopharynx smooth. Antenna 3 (one specimen)
0:35 mm long, 0-18 length of wing, 1:29 length of 4+. 5, 2:10 length of labrum; two ascoids on segments
3-8 (rest missing), that on 4 = 0-55 of its length; no papilla on 5, but several on 6 and 8 near middle.
Maxilla with six lateral teeth, three of them very small, and 27 ventral teeth, the proximal ones very small;
dental depth 0:09 mm; palpal ratio 10 : 28 : 40 : 34 : 62. Scutum brown and pleura pale. Inter-precoxal
lobes narrow. Wing length 1-96 (1-89-2-03) mm, about 3-1 times width, R,/R2,3; 2:36 (1:96-2:88), R,
overlap/R, 0-73 (0:67-0:79). Leg ratios: fore (0-69), 10:11-2:—:mid (0-69), 10: 13-9:0-75; hind
(0-77), 10: 15-5: —-. Spermatheca thin-walled and oblong.
Discussion. The specimens described here differ from S. dapsilidentes in the shape of the pig-
ment patch, weak chitinous arch and shorter palp 4, and from S. mangana in the longer antenna 3.
They differ from the description of S. torrechantei in having more cibarial teeth and a different
pigment patch, and palpal segment 3 = 1-44 times as long as 2 (instead of 1-13 times). How-
ever, the dark cibarial walls tally, and palpal segments are somewhat flexible. It seems advisable
to regard these specimens as S. torrechantei, at least till more examples of this species are found,
rather than to create a new species on slender evidence.
MATERIAL EXAMINED.
Philippines: Luzon, Nueva Viscaya, 10.iv.1968 (M. D. Delfinado & D. E. Hardy), 4 9.
DISTRIBUTION. Philippines: Apali, Malinao, Sipocot (Quate, 1965: 28; Quate & Rosario, 1962 : 797);
Nueva Viscaya (as above).
Sergentomyia (Parrotomyia) yoshimotoi (Quate)
(Map 8)
Phlebotomus (Sergentomyia) yoshimotoi Quate, 1965: 24. Holotype 9°, PHILIPPINES (BPBM) [not ex-
amined ].
In this small species with a brown scutum the female has 14—18 short cibarial teeth in a convex
row, no pigment patch and pharyngeal teeth.
DisTRIBUTION. Philippines: San Francisco area (Quate, 1965 : 26).
267
Subgenus GRASSOMYIA Theodor
Sergentomyia squamipleuris group Theodor, 1948 : 102.
Sergentomyia subgenus Grassomyia Theodor, 1958 : 47; Perfil’ev, 1968 : 339. Type-species: Phiebotomus
squamipleuris Newstead, 1912, by original designation.
The spermatheca and the expanded tips of the genital filaments, alone, distinguish this small
subgenus from others. Its distinctive nature led Abonnenc (1972) to equate it with Phlebotomus
and Sergentomyia. Grassomyia is very widely distributed in the Region.
Sergentomyia (Grassomyia) indica (Theodor)
(Map 8)
Phlebotomus squamipleuris Newstead ; Sinton, 19236 : 65; 1924a : 813; 1927c : 947; 1927d : 27; 1927e : 31;
1928c : 321; 1932a: 60; 1933e:418, 422; Perfil’ev, 1939: 82 [in part]; Mitra & Roy, 1955a: 324
[abdominal hairs].
Phlebotomus squamipleuris var. indicus Theodor, 1931 : 470; 1938 : 269; Lewis, 1957 : 168. Type, ‘INDIA’
(TC) [not examined].
Phlebotomus (Prophlebotomus) sauamipleuris var. indicus Theodor; Parrot, 1937 : 119; Parrot & Clastrier,
1952 : 164.
Phlebotomus squamipleuris var. poonaensis Mitra & Mitra, 1953 : 434 [hairs]; Mitra & Roy, 1954: 191
[description]. Syntypes 2, INDIA (Museum of the Armed Forces Medical College, Poona) [not ex-
amined]. [Synonymized by Abonnenc, 1967 : 115; 1969 : 308.]
Phlebotomus (Sergentomyia) squamipleuris indicus Theodor; Quate, 1962b : 259 [variation].
Phlebotomus (Grassomyia) squamipleuris indicus Theodor; Abonnenc, 1967 : 114.
Sergentomyia (Grassomyia) squamipleuris indica (Theodor); Lewis, 1967 : 34.
Sergentomyia (Grassomyia) indica (Theodor); Theodor & Mesghali, 1964 : 296 [India & China]; Perfil’ev,
1968 : 344; Lewis & Dyce, 1976: 210 [Bali form].
The combination of a convex cibarial tooth-row and rounded spermatheca, in the female, and
filaments with dilated tips, in the male, distinguishes this from all other Oriental species.
Various forms of Grassomyia in different parts of the Old World have undergone nomen-
clatorial changes, some of which are as follows. S. squamipleuris (Newstead) was described from
the Sudan in 1912, S. ghesquierei (Parrot) from Zaire in 1929, S. s. indica in 1931, S. dreyfussi
(Parrot) from Algeria in 1933, S. inermis (Theodor) from Africa in 1938, and several eventual
synonyms at various times. Perfil’ev (1939) placed all members of the complex in one species,
without individual names. Quate (19625 : 260) treated indica as a synonym of squamipleuris and
(1964) regarded dreyfussi, inermis and ghesquierei also as synonyms. Theodor & Mesghali (1964)
recognized four species, S. dreyfussi, indica, inermis and squamipleuris, and described S. d.
turkestanica from Turkestan and eastern Iran. Perfil’ev (1966) recognized five species and
described S. s. karakalensis from Soviet Central Asia, and (1968, in letter) informed me that this
was a synonym of turkestanica which might be treated as a subspecies of squamipleuris. S. mada-
gascarensis (Abonnenc) was described in 1969. O. Theodor (1971, in letter) informed me that
squamipleuris from Taiwan differed distinctly from indica, the pharynx being narrower, pigment
patch of different form and cibarial teeth only about 25 in number, and he considered it as good
a species as the African ones recognized by Abonnenc in 1969. Lewis (19745) regarded the Yemen
form as S. dreyfussi, and Lewis & Dyce (1976) studied the Bali form of S. indica which differed
from West Malaysian and Indian specimens.
According to Theodor and Mesghali, the females of indica and turkestanica, its nearest geo-
graphical relative, have respectively: antenna 3/labrum 0-9 and 1-1—-1-2; cibarial teeth, 33-36
and 40; antenna 3 length, 0-12-0-17 and 0-2-0-22 mm;; spines on femur 1, none and three to four.
Ten females from Peshawar were examined and had antenna 3/labrum of 0-93 (0-91-0-97).
Cibarial teeth were difficult to count in uncrushed specimens, but one chosen at random and
so treated showed 46 teeth. Antenna 3 was 0-14 (0:14-0:15). Femoral spines are often lost and
their sockets look like those of large hairs. None was seen on many flies examined.
The divergence of views, variation in the complex and appearance of new forms from time to
time make it difficult to classify some. For the present I am treating the Oriental form as a species,
268
——
S. indica, in which the number of teeth may be unimportant but other characters may later indi-
cate the need for subdivision. The mesanepimeron is bare, as in two African forms.
DISTRIBUTION. Cambodia: Phnom Penh (Parrot & Clastrier, 1952:153). China: Hainan (Leng,
1964 : 127); Aihsien, Lingshui, Paoting (Yao & Wu, 1940: 797; 1941b:77). Hong Kong: Saikung
(W. J. Voss & Hui Wai Ming, via BPBM). India: Bhavnagar, Guntur, Itauriga, Tittagash, Undi, Vellore
(BMNH); Pusa (about as common as P. argentipes according to Craighead & Das, 1928 : 863), Ranaghat
(S. Das); Bir, Parbhani (V. Dhanda); Riasi (Jacob & Kalra, 1951 : 325); Kota (Kaul et a/., 1951 : 325);
Karnal, Saharanpur (Lewis, 1967: 35); Mendhar, Punch (Mitra, 1959: 62); Jammu, Nedumangad,
Poona, Singanama, Uri (Mitra & Roy, 1953a: 324); Kasauli, Madras, Pipariya (Sinton, 19235: 66);
Calcutta, Golaghat, Patna (Sinton, 1927b : 942); Bombay, Faizabad, Itaunia, Laharpur, Nabinagar,
Narnaul, Roorkee, Trivandrum (Sinton’s notes); Naini Tal area (Wattal et al., 1967). Laos: Luang
Prbang, Ventiane (Quate, 1962b : 259). Nepal: Pokhara (L. W. Quate via BPBM), 21 9, 4 3. Nusa Teng-
gara: Dili, Kabaru, Los Palos, Same, Suai, Sumbara, Tafara Cape, Waingapu Bay (Lewis & Dyce,
1976 : 210). Pakistan: Dehra Ismail Khan, Gujrat, Jhelum, Khanki, Lahore, Peshawar, Rawalpindi,
Said Pur, Saidu Sharif, Taxla (Lewis, 1967 : 37); Cherat, Shikarpur, Tank (Sinton’s notes). Taiwan:
Henghun area, Pingtung area, Taitung (lowlands south of 32° north, Cates & Lien, 1970 : 535). Thailand:
Ayutthaya, Bangkok, Chieng Mai, Meung district, Phra Phutthabat district, Tha Li district (Quate,
19625 : 259); in trains (Sinton, 1931d: 104). West Malaysia: Rantau Panjang (Lewis, 1957 : 168); Carey
Island (A. Rudnick).
In India S. indica is widely distributed all over the plains and up to 1830 m (Sinton, 1932a).
Subgenus NEOPHLEBOTOMUS Franca & Parrot
Phlebotomus subgenus Neophlebotomus Franca & Parrot, 1920 : 699. Type-species: Phlebotomus malabari-
cus Annandale, 1910, by original designation.
Sergentomyia zeylanica group Theodor, 1948 : 101.
Sergentomyia subgenus Rondanomyia Theodor, 1958 : 48; Perfil’ev, 1968 : 335. Type-species: Phlebotomus
malabaricus Annandale, 1910, by original designation. Syn. n.
S. malabarica is now seen to belong in ‘Rondanomyia’, but is the type-species of Neophlebotomus
(discussed by Theodor, 1948 : 88, 89; Kirk & Lewis, 1951 : 406) which was regarded as a syno-
nym of genus Sergentomyia. The original characterization of Neophlebotomus was indefinite and
tentative, and was based on a non-existent character, but it is technically valid nevertheless
(ICZN, 1964: Articles 17(8) and 18(a)). It is therefore treated here as a subgenus, with Ron-
danomyia as a synonym.
A long antenna 3, a broad wing with long R, and a style with two spines near the middle class
some species as members of this subgenus, but all the features mentioned in couplet 110 of the
key must often be considered, and certain species are difficult to classify subgenerically. The
figure for R,/R,,3 in six Palaearctic species studied by Theodor (1958) is 1-30 (0-5-1-98), but if
the exceptional S. pawlowskyi Perfil’ev is omitted the figure is 1-7 (0-75-1-98).
Species of the subgenus occur mainly in areas of high rainfall in the east of the Region where
they are comparable with central African members of the subgenus.
Some species of Sergentomyia were placed by Theodor (1948) in the zeylanica group which
was divided into two series according to the nature of the style. Theodor (1958) converted the
group to the subgenus Rondanomyia but did not mention the two series.
The spermatheca is of some use as a subgeneric character but, as in some other groups, is often
difficult to see and define. Although Theodor refers to the tubular spermatheca, it may sometimes
be elliptical (Theodor, 1938 : 264).
The species can be grouped (Table 1) according to cibarial armature of the females and what
is known of the spermatheca, sometimes from imperfect specimens.
In the dhandai series the spermatheca is long and cylindrical.
In the arboris series the spermatheca is more or less oblong, with the knob in a pit, and the
female has three or more rows of cibarial fore teeth. S. arboris is closely related to S. gombaki,
and S. malayae to S. zeylanica, and further knowledge of their distribution and variation is
needed, with special reference to the length of the labrum.
269
In the balica series the spermatheca is like that of the arboris series and the cibarium of the
female has less than three rows of fore teeth. S. jefferyi differs from most sandflies in its light
maxillary armature.
In the quatei series the spermathecal knob protrudes proximally and there is no collar. S. traubi
has a comparable but short spermatheca.
In S. hamidi the spermathecal duct starts as a funnel.
This arrangement, which is based on females partly for their biological interest, cuts across
Theodor’s (1948) sections a and b (based on the central or distal position of two of the spines on
the style of the male).
Sergentomyia (Neophlebotomus) arboris (Sinton)
(Map 8)
Phlebotomus arboris Sinton, 193le : 107; Theodor, 1938 : 263 [2, previously mistaken for S. zeylanica];
Parrot, 1940 : 311; 1946: 71. About 20 syntype 3, INpIA (depository unknown) [not examined].
Sergentomyia arboris (Sinton); Theodor, 1948 : 111.
Sergentomyia (Rondanomyia) arboris (Sinton); Lewis, 1973a : 250.
The presence of about eight rows of cibarial fore teeth and the lack of a definite forward process
of the pigment patch differentiate the female from all other species.
2 (extra facts). Labrum 0-32 mm long, 0-14 length of wing (2:26 mm). Hypopharynx with smooth tip and
low undulations. Antenna 3 = 0:39 mm long, 0-17 length of wing. Maxilla with eight lateral and 58
ventral teeth, dental depth 0-14 mm.
MATERIAL EXAMINED.
India: Baragi, 24.xii.1957 (H. Trapido), 1 9.
DISTRIBUTION. India: Marianbari (Sinton, 193le: 107); Darjeeling, (Theodor, 1938 : 263); Baragi,
Hisur, Kannur (H. Trapido). Sri Lanka: Depanama, Katuwala (Theodor, 1938 : 264).
Sergentomyia (Neophlebotomus) balica Lewis & Dyce
(Figs 90-92, Map 8)
Sergentomyia (Rondanomyia) balica Lewis & Dyce, 1976:212. Holotype 2, SumBAWA (Australian
National Insect Collection) [examined ].
The female has 18-22 cibarial teeth in a straight line and no pigment patch. S. bukidnonis, S.
coronata (Quate & Quate), S. crypta (Quate & Quate), S. curtata (Quate) and S. sansaporensis
Fairchild have a rather similar cibarium but a relatively shorter antenna 3. The male is described
below for the first time.
The following measurements of five females from Java (Djakarta, xi.1908; Semarang, i.1910, two;
ii.1910, two) agree well with Bali specimens. Labrum 0-20 (0:19-0:20) mm long, 0-10 (0:10-0:10) length
of wing. Cibarium with 16-20 hind teeth, accessory fore teeth usually present. Antenna 3 = 0-33
(0:31-0:34) mm long, 0-17 (0:16-0:18) length of wing, 1-36 (1-:32-1-44) length of 4+5, 1-67 (1-59-1-74)
length of labrum. Palpal ratio about 10: 23 : 38: 48: 67. Wing length 1-97 (1-94-1-98) mm, 3-4 times
width, R./R2.3 0°83 (0°69-0-93). R, overlap/R, 0-53 (0-49-0-57). Spermathecal ducts not clearly visible.
One incomplete female from Djakarta, xi.1908, apparently of a form near balica, has eight cibarial hind
tooth and no fore teeth. The labrum is 0:17 mm long and 0-10 length of wing. Antenna 3 is 0-35 mm long,
0-21 length of wing and 2:12 length of labrum. The wing length is 1-68 mm, about 3-5 times width,
R,/ R23 0:77, and R, overlap/R, 0-47.
$. Labrum 0-17 (0:17-0:18) mm long, 0:10 (0:10-0:11) length of wing. Cibarium with about eight
teeth, some with several points, two or three specks representing fore teeth, and no pigment patch.
Pharynx almost unarmed. Antenna 3 = 0-35 (0:35-0:37) mm long, 0-21 (0-20-0-21) length of wing,
1-34 (1:31-1:39) length of 4+5, 2-01 (1:89-2:10) length of labrum, one ascoid on segments 3-14 (15
lacking), that on 4 being about 0:28 length of segment. Wing length 1-70 (1:66-1:74) mm, 3-5 times
270
width, R./ R23 0°70 (0:66-0:74), R, overlap/R, 0:35 (0:32-0:37). Aedeagus tapering and bluntly pointed.
Paramere beaked. Coxite with about 25 scattered brush hairs, mainly in distal half. Style with seta at
0:54, two spines subterminal and two terminal.
DISTRIBUTION. Borneo: Sabah, Tawau (L. W. Quate, a female with 24 hind teeth and two rows, of four
and eight, fore teeth, which may be this species). Java: Djakarta, Semarang (E. Jacobson). Nusa Tenggara:
Denpasar area, Pedang Bay, Sumbawa (Lewis & Dyce, 1976: 212). West Malaysia: Kuala Lumpur
(5.vii.1923, H. M. Pendlebury, 1 9); Carey Island (attracted to man or to CO,-light-trap, 2 9, 3 3), Port
Dixon (1 9) (A. Rudnick).
Sergentomyia (Neophlebotomus) sp. (Besout)
(Figs 93-94, Map 8)
This species differs from many by having some 95 hairs in the coxite brush. It is left without a
formal name till females can be associated with the males.
36. Labrum 0-20 (0:20-0:21) mm long, 0-13 (0-12-0-13) length of wing. Cibarial hind teeth in the form of
minute spicules, roughly arranged in about 12 groups on a concave arc; fore teeth comprising a row of
about 12. Pharynx unarmed. Antenna 3 = 0-33 (0:30-0:36) mm long, 0-21 (0:20-0:21) length of wing,
1.37 (1:27-1:34) length of 4+ 5, 1-63 (1:54-1-72) length of labrum; one ascoid on segments 3-15, that on
4 = 0°55 length of segment. Maxilla with about 20 vestigial ventral teeth. Palpal segment 4 = about
1-3 times length of 3. Scutum pale brown and pleura very pale. Wing length 1-60 (1:51—1-70) mm, 3 times
width R,/R2,5 1:20 (1-:10-1-30), R, overlap/R, 0:54 (0-53-0-61). Aedeagus very pale except at base.
Paramere with beaked and swollen tip. Coxite with discrete dense patch of about 95 stout hairs. Style
broad at centre, with a small seta at about 0-37, setae at about 0-44 and 0-6, and two at tip.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, 19.x.1973 and 15.vi.1974 (J. Jeffery), 3 3; Punjong
Rabok, 17 and 27.xi.1968 (A. Rudnick).
Sergentomyia (Neophlebotomus) chakravarti (Mitra) comb. n.
(Map 8)
Phlebotomus chakravarti Mitra, 1953b : 158. Holotype 9, InpIA (Armed Forces Medical College, Poona,
apparently lost, according to V. Dhanda, 1970, in letter) [not examined].
The female has about 14 cibarial teeth on a line angular at the centre, and seven round teeth
behind the hind ones.
DISTRIBUTION. India: Hoora (Mitra, 19535 : 158).
Sergentomyia (Neophlebotomus) dhandai Lewis sp. n.
(Figs 95-101, Map 8)
This species differs from S. hodgsoni in having more cibarial hind teeth.
9. Labrum 0-22 (0:21-0:23) mm long, 0-10 (0:10-0:11) length of wing. Cibarium with about 24 con-
tiguous pointed teeth on a deep arc, and a broad pigment patch, cuticle at base of teeth very dark, and
chitinous arch inapparent; salivary pump wide. Pharynx with faintly spiculate ridges. Hypopharynx with
low undulations. Antenna 3 = 0-27 (0:24-0:29) mm long, 0-13 (0-11-0-14) length of wing, 1-13 (1:03-1:23)
length of 4+5, 1-21 (1:15-1:28) length of labrum, two ascoids on segments 3-15, that on 4 = 0°5 of its
length. Maxilla with 11 lateral and 36 short broad ventral teeth, dental depth 0-11 mm; palpal ratio
10: 21 : 35 : 37: 85, segment 3 with clavate sensilla bunched on basal half. Scutum and much of pleura
dark. Wing length 2-13 (2:11-2:15) mm, 3:3 times width, R,/R2,3 1-40 (1:27-1:53), R, overlap/R,
0-58 (0:58-0:58). Femur 1 with six large sockets and at least three spines, femur 2 with five large sockets.
Hind leg in holotype, lengths in mm: femur 0-44; tibia, 0-57; basitarsus, 0-27; whole, 1:68. Spermatheca
271
OL 1 ee a |
ak
Figs 93-105 Sergentomyia species. 93, 94, S. Besout sp. 3, cibarium and terminalia. 95-101, S.
dhandai: (95-99) °, cibarium, pharynx, maxilla and spermatheca; (100, 101) 3, cibarium and
terminalia. 102-105, S. gemmea, 2: (102, 103) labrocibarium and tip of labrum; (104) cibarium;
(105) pharynx.
272
subcylindrical with slightly expanded tip, thick walls and faint internal wrinkles which are sometimes
invisible; knob short and cylindrical; ducts faintly striated and joining a common duct.
$. Labrum 0-18 (0:18-0:18) mm long, 0-11 (0:10-0:11) length of wing. Cibarium with about 24 pointed
teeth on a deep arc, no pigment patch. Pharynx almost unarmed. Antenna 3 = 0-28 (0-27-0-29) mm
long, 0-16 (0-15-0-17) length of wing, 1-11 (1-11-1-11) length of 4+.5, 1-54 (1-46-1-63) length of labrum,
one ascoid on segments 3-14 (15 lacking), that on 4 about 0-36 of its length. Wing length 1-72 (1-:71-1-74)
mm, 3-7 times width, R/.R,,3 1.41 (1:21-1:62), R, overlap/R, 0-55 (0:51-0:59). Femur 1 with four spines,
femur 2 with at least one. Aedeagus with rounded bluntly pointed tip. Filament 4:5 times length of pump.
Style with two of spines subterminal at different levels, and seta at 0-6.
ComMENTS. S. dhandai apparently differs from S. chakravarti, from the same general area, in
having more cibarial teeth, a different pigment patch, and no teeth behind the main row. Such
differences could conceivably be due to different mounting methods, but the shorter leg of S.
dhandai in a rather larger fly (with longer antenna 3 and wing) indicates that they are different
species.
MATERIAL EXAMINED.
Holotype 9, India: Poona district, Bhor Ghat, 27.ix.1969 (S. N. Guttikar, G. B. Modi & P. V. M. Mahadev),
tree hole in hilly woodland (BMNH).
Paratypes. India: same data, 1 2, 2 ¢ (BMNH). Other specimens, examined by Dr V. Dhanda, are in the
Virus Research Centre, Poona.
Sergentomyia (Neophlebotomus) gemmea Lewis & Jeffery sp. n.
(Figs 102-113, Map 9)
The large size of some of the cibarial fore teeth characterizes this species. The name, meaning
jewelled, indicates their ornamental appearance.
9. Labrum 0-32 (0:29-0:34) mm long, 0-16 (0:15-0:17) length of wing, apical sensilla normal, adorals
small. Cibarium with ten hind teeth with broad bases narrowing abruptly to fine points, with one row of
eight very large fore teeth, two rows of small teeth in front of them, and a patch of small fore teeth at each
side; pigment patch pale; arch strong. Pharynx with linear and with finely spiculate ridges. Hypopharynx
with distinct teeth wider than high. Antenna 3 = 0-36 (0:34-0:39) mm long, 0-18 (0:18-0:20) length of
wing, 1.26 (1:22-1:31) length of 4+5, 1-13 (1:08-1:19) length of labrum; two ascoids on segments
3-15, that on 4 = 0-87 length of segment and reaching next one, with spur; no papilla on 5. Mandible
pointed. Maxilla with about eight lateral teeth and about 41 ventrals, a few of them very small, dental
depth 0-12 mm, palpal ratio 10: 16: 31:34:61. Scutum pale brown and pleura mainly pale. Wing
length 1-95 (1:98-2:00) mm, 2-7 times width, R./Rj.3 1:41 (0:99-1:72), R, overlap/R, rather constant
near 0-61 (0:52-0:68). Abdominal tergites 3-6 with a few erect hairs on hind margins. Spermatheca
narrow with some wrinkles proximally, with knob in deep narrow pit, delicate ducts uniting into short
common one.
$. Labrum 0-22 mm long, 0-13 length of wing. Cibarium with about six irregular hind teeth and about
20 irregular fore teeth of which a few posterior ones are slightly larger than the others; pigment patch
indefinite. Pharynx with faint ridges. Antenna 3 = 0:35 mm long, 0:21 length of wing, 1:21 length of
4+5, 1-59 length of labrum, one ascoid on segments 3-15, that on 4 0:64 length of segment; no papilla on
5. Wing length 1-64 mm, 3 times width, R./R:,3 1:01, R, overlap/R, 0-19. Genital filament 3-1 times length
of pump. Coxite broad with patch of about 43 narrow hairs merging into large meso-dorsal hairs; style
- with seta at 0-65 and two of stout spines at about 0-68.
MATERIAL EXAMINED.
Holotype 9, West Malaysia: Gunong Besout Forest Reserve, 20.iii.1974 (A. B. Knudsen & colleagues)
(BMNH).
Paratypes. West Malaysia: Pulau Meranti, 19.xi.1970, 11.ii.1971 (S$. Mahadevan), trap baited with
monkey Macaca nemestrina (beroks) in swamp forest canopy 8-5 m above ground, 19-00-21.00 hours,
2 2; as for holotype, 22.ii.1974 (J. Jeffery & colleagues), 1 2, 1 3; Klang, 1.iii.1972 (D. J. Lewis), 1 2;
Tanjong Rabok, 12.xi.1969 (A. Rudnick), 1 9. (Allin BMNH.)
273
a er er ae a
=e ane ae o oe
2075 (ee ey See ay)
a
=
° oz
106:1°*.,
os og
108
Figs 106-118 Sergentomyia species. 106-113, S. gemmea: (106, 107) 2, antenna 4; (108) 92, maxilla;
(109) 9, wing; (110, 111) 9, spermathecae; (112, 113) 3, cibarium and terminalia. 114-118, S.
gombaki, 2: (114) head; (115, 116) labrocibarium and tip of labrum; (117, 118) sagittal section of
fascicle and cibarium.
274
Sergentomyia (Neophlebotomus) gombaki (Lewis & Wharton) stat. n.
(Figs 114-126, Map 9)
Phlebotomus sp. Lewis, 1957 : 167.
Phlebotomus (Sergentomyia) zeylanica gombaki Lewis & Wharton, 1963 : 121. Holotype 9, West MALAYSIA
(BMNH) [examined].
Sergentomyia (Rondanomyia) zeylanica gombaki (Lewis & Wharton) Lewis, 1973a : 251.
The cibarium of the female has about eight rows of fore teeth and the process of the pigment
patch is faint or absent. The labrum is so long that the species looks superficially rather like a
Phlebotomus. The male is described for the first time.
2. Labrum 0-40 (0:34-0:42) mm long, 0-20 (0:18-0:21) length of wing, first and last apical sensilla pro-
minent and adorals small. Cibarium with 12 pointed teeth and about seven rows of fore teeth; pigment
patch very large and rounded anteriorly. Pharynx with faint ridges. Hypopharynx with low teeth and a
smooth tip. Antenna 3 = 0-35 (0:31-0:38) mm long, 0-17 (0-:16-0-18) length of wing, 1-26 (1-20-1-34)
times length of 4+ 5, 0-88 (0:83-0:90) length of labrum, two ascoids on 3-15, that on 4 = 0-7 length of
segment, with a spur, no papilla on antenna 5. Mandible with about 130 teeth including proximal faint
ones, pointed. Maxilla with seven broad lateral teeth and 70 ventrals, dental depth 0:21 mm; palp delicate
and difficult to measure, approximate ratio 10 : 20 : 22 : 23 : 57. Scutum brown and pleura mainly pale.
Wing length 2-01 (1:97-2:09) mm, 3-1 times width, R./Ro,3 1:27 (1:07-1:62), R, overlap/R, 0-64
(0:57-0:69). Spermatheca oblong with no collar, and ductules entering deep pit.
$. Labrum 0:27 (0:25-0:29) mm, 0-14 (0:14-0:15) length of wing. Cibarium with about 16 hind teeth
in groups of two or three, and about 16 scattered fore teeth. Pharynx with faint ridges. Antenna 3 = 0-44
(0:41-0:46) mm long, 0-23 (0:21-0:25) length of wing, 1-15 (1:09-1:19) times length of 4+5, 1-62
(1:53-1:70) times length of labrum, one ascoid on segments 3-15, that on 4 = 0-4 of its length. Palpal
ratio 10: 35:42:42: 74. Wing length 1-92 (1:83-1:97) mm., 3-6 times width, R,/R.,3; 0-79 (0-48-1 -09),
R, overlap/R, 0-52 (0:38-0:61). Aedeagus pointed. Paramere beaked. Coxite delicate with sub-basal
dense patch of about 183 hairs. Style with two spines terminal and two at 0-6, and a seta at 0°5.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, April 1973 to August 1974 (A. B. Knudsen & colleagues),
294 2, 218 3, 10 of each measured.
DISTRIBUTION. West Malaysia: Bukit Ibam (L. W. Quate, via BPBM); Gunong Besout area (as above);
Ulu Gombak (Lewis & Wharton, 1963 : 121); Batang Padang, 3.vi.1923, 700 m (H. M. Pendlebury).
Sergentomyia (Neophlebotomus) hamidi Lewis & Jeffery sp. n.
(Figs 127-131, Map 9)
The female has a cibarium rather like those of S. quatei and S. silvatica, but different sperma-
thecae.
9. Eye 0-67 length of head. Labrum 0-16 (0:15-0:17) mm long, 0-09 (0-08-0-09) length of wing, adoral
sensilla large. Cibarium with about two to four main spiky teeth, more or less continuous with a group of
about ten small teeth at each side, pigment patch elongated and pale. Pharynx with a few lines and
spicules. Hypopharynx with low undulations. Antenna 3 = 0-36 (0:35-0:36) mm long, 0-19 (0:19-0:20)
length of wing, 0-78 (0:74-0:82) length of 4+5, 2:25 (2:19-2:29) length of labrum, two delicate ascoids,
difficult to see, on segments 3-15, that on 4 being 0-28 length of segment, no papilla on 5. Mandible
rather blunt, with large main teeth, 1-7 zm wide and very small distal ones. Maxilla with 12 lateral and 24
ventral teeth, dental depth 0-08 mm, palpal ratio 10 : 15 : 30 : 17 : 39. Scutum brown and pleuron brown-
ish. Wing length 1-85 (1:79-1:90) mm, about 3-4 times width, R,/Rz+3 0°85 (0:72-0:97), R, overlap/R.
0:44 (0:33-0:53). Spermatheca with faint ductules, a refractive knob, very thin walls with minute spiculi-
form specks, narrowing toward the duct which begins as a relatively thick-walled funnel.
MATERIAL EXAMINED.
Holotype 2, West Malaysia: Gunong Besout Forest Reserve, 29.v.1974 (Ahmad bin Abd. Hamid) (BMNH).
Paratypes. Same locality, 18.iii.1974 (A. B. Knudsen et al.), 1 2; 13.vi.1974 (J. Jeffery et al.), 1 9; 1974
(R. B. Tesh), 2 2. (Two in BMNH, two in U.S. National Museum.)
Non-paratypic material. West Malaysia: Bukit Ibam (L. W. Quate, via BPBM).
275
AWelkn,
Figs 119-133 Sergentomyia species. 119-126, S. gombaki: (119) 9, cibarium and base of pharynx;
(120) 2, pharynx; (121) 2, antenna 4; (122) 9, maxilla; (123) 9, wing; (124) 2, tip of spermatheca;
(125, 126) 3, terminalia and pump. 127-131, S. hamidi, 2: (127, 128) labrocibarium and cibarium;
(129-131) spermatheca and ends. 132, 133, S. iyengari, 2, labrocibarium and tip of labrum.
276
wah 6
_ =
ata».
Sergentomyia (Neophlebotomus) hodgsoni (Sinton)
Phlebotomus hodgsoni Sinton, 1933a : 874.
The cibarium of the female has 42-68 teeth, and the paramere of the male bears a spinose pro-
cess. The species occurs from Central Asia to western India.
Sergentomyia (Neophlebotomus) hodgsoni hodgsoni (Sinton)
(Map 9)
Phlebotomus hodgsoni Sinton, 1933a : 874; 1933d : 226; 1933e : 419. Lectotype 2, PAKISTAN (BMNH), de-
signated by Lewis, 1967 : 37 [examined].
Sergentomyia (Rondanomyia) hodgsoni (Sinton); Theodor & Mesghali, 1964 : 296.
Sergentomyia (Rondanomyia) pawlowskyi hodgsoni (Sinton); Lewis, 1967 : 37.
Sergentomyia (Rondanomyia) hodgsoni hodgsoni Artemiev, 1976a : 39.
The cibarium of the female has 50-60 teeth, and the paramere of the male bears a ventral spinose
process.
DISTRIBUTION. India: Mahasu area (Rao et al., 1973). Pakistan: Cherat, Gwadi, Jandola, Landi Kotal,
Parkuta, Peshawar, Rawalpindi, Said Pur, Taxla (Lewis, 1967 : 38).
Sergentomyia (Neophlebotomus) iyengari (Sinton) comb. n.
(Figs 132, 133, Map 9)
Phlebotomus iyengari Sinton, 1932c : 221; Raynal & Gaschen, 1935c : 507 [2]; Raynal, 19355 : 249, 294;
Theodor, 19384: 268; 19385: 172 [S. dureni (Parrot) of Africa as possible variety]. Syntypes 2 9,
INDIA (depository unknown) [not examined].
Phlebotomus hivernus Raynal & Gaschen, 1935d : 582. Syntypes 4 9, 4 3, VIETNAM (NorTH) (depository
unknown) [not examined]. [Synonymized by Quate, 19625 : 265.]
Phlebotomus hibernus Anonymous, 1935 :779 [unjustified emendation under ICZN Article 33a(ii)];
Raynal, 19355 : 236, 239; Parrot, 1940: 311.
Phlebotomus iyengari var. hivernus (Raynal & Gaschen) Theodor, 1938 : 267.
Phlebotomus iyengari var. malayensis Theodor, 1938 : 266. Syntypes 5 2, 173, WEST MALAysIA (depository
unknown) [not examined]. [Synonymized by Quate, 19626 : 265.]
Phlebotomus (Prophlebotomus) hibernus Anonymous; Parrot, 1940 : 311.
Phlebotomus iyengari var. hainanensis Yao & Wu, 1940 : 786; 1941 : 77; Leng, 1964: 121, 128. Syntypes
3 9, CHINA (depository unknown) [not examined]. [Synonymized by Quate, 19625 : 265, 266. ]
Phlebotomus (Prophlebotomus) iyengari Sinton; Parrot, 1946 : 70; Parrot & Clastrier, 1952 : 159.
Phlebotomus (Sergentomyia) iyengari var. malayensis Theodor; Lewis, 1957 : 168.
Phlebotomus (Sergentomyia) iyengari Sinton; Quate, 19626 : 265.
Phlebotomus (Sergentomyia) iyengari taiwanensis Cates & Lien, 1970 : 530. Holotype 9, TAIWAN (Taiwan
Malaria Research Institute) [not examined]. Syn. n.
Sergentomyia (Rondanomyia) iyengari (Sinton); Lewis, 1973a : 250; Yen-Chia, 1977 : 336.
The cibarium of the female has the central teeth smaller than the rest, fore teeth absent or ranging
from one row of four to two rows of up to 20, and a pigment patch with forward projection thick,
small or absent.
Theodor (1938) pointed out that this species is related to S. dureni (Parrot) of Africa. S. iyen-
gari, like S. tambori, somewhat resembles the African S. decipiens (Theodor) and S. dureni but
has a very pronounced spermathecal collar.
In a female from Gunong Besout, the labral apical sensilla are clearly visible and there are few
anterior adorals, the hypopharynx has a few very low serrations, the mandible is normal, and
the maxilla has eight broad lateral teeth, 26 well-marked ventrals and a dental depth of 0-10 mm.
The extra ascoids in the male, mentioned by Parrot (1940), are narrow with small bases and
resemble colourless spines near the tips of segments 8-16 and on antennae of some other species.
They are perhaps best regarded as vestiges to be excluded from the antennal formula.
2k
The form taiwanensis is treated here as a synonym in view of Quate’s conclusions on variation
in the species. Yen-Chia (1977) recognizes S. hainanensis as a species with malayensis as its
synonym.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, 1973-74 (A. B. Knudsen, J. Jeffery & colleagues), 24 9,
18 3.
DIsTRIBUTION. China: Hainan Island (Leng, 1964: 127); Aihsien (Yao & Wu, 1940: 797); Lingshui,
Paoting (Yao & Wu, 19416 : 77). India: Trivandrum (Sinton, 1933e : 221). Laos: Luang-Prabang (Parrot
& Clastrier, 1952 : 153); Muong Sing (Quate, 19625 : 265). Taiwan: many lowland places except west
coastal plain, Hua-lien area, Kao-hsiung area, Taipei area (Cates & Lien, 1970 : 531). Thailand: Chieng
Mai, Doi Sutep, Khon Kaen Province, Nong Khai area, Sara Buri, Tha Li area, Udon Thani area
(Quate, 19625 : 265). Vietnam (North): Bim Son, Cho Ganh (Raynal, 19355 : 253, 297). West Malaysia:
Gunong Besout area (as above); Kuala Lumpur area, Ulu Gombak (Lewis, 1957 : 168); Selangor State
(Theodor, 1938 : 267).
In Indo-China Raynal (1936a : 361) at first found S. iyengari in only one locality (occasionally
at the start of the warm season), and in the north (1936a : 367) encountered it only on mountain
spurs about 20° north in the cold season.
Sergentomyia (Neophlebotomus) jefferyi Lewis sp. n.
(Figs 134-142, Map 9)
The female differs from S. hitchensi in having a prominent pigment patch.
2. Eye 0-55 length of head. Labrum 0-19 (0:17-0:20) long, 0-11 (0:11-0:13) length of wing, apical sensilla
very delicate, and subapicals hardly visible, adorals reduced. Cibarium with seven or eight delicate hind
teeth and about ten fore teeth in single row; pigment patch about half internal width of cibarium, broader
in front than behind, reddish brown, a faint short broad patch in front of it in ventral wall. Pharynx
with faint lines. Hypopharynx with very low teeth which approach the tip. Antenna 3 = 0-25 (0:24-0:28)
mm jong, 0-15 (0:15-0:16) length of wing, 1-18 (1-10-1-20) length of 4+5, 1-36 (1-32-1-45) length of
labrum, two ascoids on segments 3-15, that on 4 very thin and about 0-8 length of segment, no papilla
on 5. Mandible pointed. Maxillary teeth vestigial; palpal ratio 10 : 20 : 31 : 50: 74. Scutum pale brown
and pleura pale. Wing length 1-69 (1:58-1:75) mm, 3-3 times width, R,/R.,3 0:96 (0:74-1:15), R, over-
lap/R, 0-46 (0:29-0:57). Spermatheca usually wrinkled, with a rather low knob in a deep pit.
$. Labrum 0-16 (0:15-0:17) mm long, 0-11 (0:11-0:12) length of wing. Cibarium with about seven
ill-defined hind teeth, and about 15 small fore teeth in irregular row. Pharynx unarmed. Antenna 3 = 0-25
(0:22-0:27) mm long, 0-17 (0:16-0:20) length of wing, 1-16 (1-01-1-23) length of 4+5, 1-55 (1-41-1-71)
length of labrum, one ascoid on segments 3-15, that on 4 about 0-33 length of segment, and small
transparent hairs on some of distal segments. Wing length 1-44 (1:38-1:50) mm, 3-6 times width, Ro/Re45
0-72 (0:53-0:85), R, overlap/R, 0:30 (0:18-0:41). Coxite and style with few hairs, seta on style small.
MATERIAL EXAMINED.
Holotype 2, West Malaysia: Perak, Gunong Besout Forest Reserve, 5.v.1974 (J. Jeffery) (BMNH).
Paratypes. Same locality, 24.ii.1974 to 10.viii.1974, 12 9, 10 3, ten of each measured. (All in BMNH.)
Sergentomyia (Neophlebotomus) khawi (Raynal)
(Map 9)
Phlebotomus khawi Raynal, 1936c : 529; 1937 : 72; Yao & Wu, 19415 : 77. Syntypes 2 3, CHINA (depository
unknown) [not examined].
Phlebotomus (Prophlebotomus) khawi Raynal; Parrot & Clastrier, 1952 : 160.
Sergentomyia (Rondanomyia) khawi (Raynal); Theodor, 1958 : 48; Lewis, 1973a: 251.
Phlebotomus (Sergentomyia) khawi Raynal; Quate, 1962b : 265.
The cibarium of the female has no pigment patch and a row of contiguous teeth, convex at the
centre.
DISTRIBUTION. Cambodia: Phnom Penh (1 3, Parrot & Clastrier, 1952 : 133).
278
esd “02 5
137
139
141
144
146
ry)
Figs 134-146 Sergentomyia species. 134-142, S. jefferyi: (134-136) 9, labrocibarium, tip of labrum
and cibarium; (137) 2, pharynx; (138) 9, antenna 4; (139) 2, maxilla; (140, 141) 2, spermatheca;
(142) 3, terminalia. 143-146, S. linearis, 2: (143, 144) cibarium and pharynx; (145) wing;
(146) spermatheca.
279
Sergentomyia (Neophlebotomus) linearis Lewis sp. n.
(Figs 143-148, Map 9)
A long narrow pigment patch in the cibarium distinguishes females and males from those of
most other species of Sergentomyia.
9. Labrum 0-25 (0:24-0:26) mm long, 0-12 (0:12-0:12) length of wing. Cibarium with about 10-12
pointed teeth in nearly straight row, and no fore teeth; pigment patch narrow and linear. Pharynx with
faint ridges. Hypopharynx smooth. Antenna 3 = 0-34 (0:32-0:36) mm long, 0-16 (0:16-0:17) length of
wing, 1:26 (1-:21-1-32) length of 4+5, 1-36 (1-31-1-40) length of labrum; two ascoids on segments 3-15,
that on 4 = 0:73 length of segment; no papilla on 5. Maxilla with no lateral teeth and vestigial ventrals;
palpal ratio 10: 24: 32: 39. Wing length 2-06 (2:00-2:19) mm, about 3-3 times width, R, (very long)/
R43 2°40 (2:16-2-66), R, overlap (consistently very long)/R, 0-75 (0:73-0:76). Spermatheca subcylindrical,
with collar and delicate duct.
$3. Labrum 0-18 (0:17-0:21) mm long, 0-11 (0-10-0-13) length of wing. Cibarium with about 12 pointed
divergent teeth and no fore teeth, pigment patch linear. Pharynx almost unarmed. Antenna 3 = 0:35
(0:31-0:43) mm long, 0-20 (0:19-0:20) length of wing, 1-21 (1-07-1-30) length of 4+5, 1-89 (1-77-2-07)
length of labrum, one ascoid on segments 3-15, that on 4 = 0-55 length of segment. Wing length 1-76
(1:67-2:18), 3-5 times width, R./R,+3 2°03 (1:61-2°64), R, overlap/R, 0-73 (0-65-0-96). Genital filament
about 2:6 times length of pump. Aedeagus tapering to a blunt point. Coxite with about ten scattered hairs
in brush. Style short and narrow, with seta at 0-75 and two of spines subterminal.
The specimens were presented to the BMNH by J. A. Sinton who had labelled them Jinearis,
presumably with reference to the pigment patch.
MATERIAL EXAMINED.
Holotype 9, India: Travancore, Kolatupuzha (= Kulathurpuzha 7?) Forest Reserve, ix(?). 1934 (M. O.
T. Iyengar), tree holes (BMNH).
Paratypes. India: data as for holotype, 3 ?, 19 3; 6 9, 10 ¢ measured; Palod Forest Reserve, ix(?).1934
(M. O. T. Iyengar), 2 2, 9 3. (All in BMNH.)
Non-paratypic material. India: Kannur, ix, x.1957 (H. Trapido). (In BMNH.)
Sergentomyia (Neophlebotomus) malabarica (Annandale) comb. rev.
(Figs 149-156, Map 9)
Phlebotomus malabaricus Annandale, 1910b : 48; Brunetti, 1912 : 214; Sinton, 1924a : 833; 1924c : 1007
[$ measurements & figs]; 1928c : 321; 193le: 110; 1935e : 420, 423; Mitra, 1953 : 162. Lectotype 3,
INDIA (Zoological Survey of India), designated by Quate, 1962c : 158 [not examined].
Sergentomyia malabarica (Annandale); Theodor, 1948 : 111 [put in africana group].
Phlebotomus (Sergentomyia) malabaricus Annandale; Quate, 1962c : 158.
The cibarium of the female has about eight hind teeth in a nearly straight row.
2. Medium-sized and very dark brown. Labrum 0:20 (0:17-0:23) mm long. Cibarium with eight pointed
teeth and a distinct pigment patch. Pharynx almost unarmed. Antenna 3 = 0-28 (0:27-0:29) mm long,
1:16 (1:11-1:20) length of 4+5, 1:42 (1:25-1:64) length of labrum, ascoid on 4 about 0-56 length of
segment. Palpal ratio 10: 21 : 34: 48:90. Wing length 2-06 (1:94-2:23) mm, 3-5 times width, R2/Re+3
2°51 (2:35-2°61), R, overlap/R, 0:76 (0:74-0:80). Spermatheca tubular.
3d. Described by Sinton (1924c).
ComMENTs. S. malabarica, described when important characters were still unknown, has for
long been a problem species with regard to the structure of the male, the identity of the female,
confusion with other species, and suggested use as a type-species (see under Sergentomyia).
The male was at first stated to bear spines on the surstyle but these proved to be merely hair-
sockets (Sinton, 1924c).
Misidentification of the female has occurred (Sinton, 1927a: 933; 1927d:25, 1927e : 30;
1928c : 321; 1932a: 61. See under S. modii.).
280
yyy)
fe ee vv)
156
152
150
160 yj)
S|
SO2S. a eae eee
01
ee oe ee
-r -- Rt Rt ay a a aoa oy a
162
Sere 157
A
-05
; 147
eee
161
Ly. ee
159
Figs 147-162 Sergentomyia species. 147, 148, S. linearis, 3: (147) cibarium; (148) terminalia. 149-
156, S. malabarica, 2: (149) cibarium; (150) pharynx; (151, 152) antennal tip and segments 3 and 4;
(153) palp; (154) wing; (155, 156) spermatheca. 157-162, S. malayae, 2: (157, 158) labrocibarium
and tip of labrum; (159) hypopharynx; (160) base of ascoid on antenna 4; (161) mandible tip;
(162) maxilla. (Figs 149-156 J. A. Sinton del.)
281
The female remained unknown till 1934 when Sinton received specimens from Kerala. He left
notes in the BMNH from which the above description was prepared.
The species is placed in this subgenus on the basis of the style-spines of the male and the wing
and spermatheca of the female.
Slides presented to the BMNH by Sinton and bearing a provisional name malabarica included
two other species, S. modii, which had previously been described as the female of malabarica,
and S. kauili.
DIsTRIBUTION. India: Travancore (now in Kerala), below western slopes of Western Ghats, Maddathorai
and Palod (= Pallode ?) (Annandale, 1910: 49); Kulathurpuzha Forest Reserve and Palod Forest
Reserve, 1934 (M. O. T. Iyengar), 4 9, 4 3.
Sergentomyia (Neophlebotomus) malayae (Lewis) stat. n.
(Figs 157-164, Map 9)
Phlebotomus (Sergentomyia) zeylanica malayae Lewis, 1957 : 166; Quate & Fairchild, 1961 : 218. Holo-
type 2, WesT MALaysIA (BMNH) [examined].
Sergentomyia (Rondanomyia) zeylanica malayae; Lewis, 1973a : 251.
The female has about three rows of cibarial fore teeth and a very long labrum, 0-18-0-20 of the
wing length, and more maxillary ventral teeth than does S. zeylanica.
9. Labrum 0-50 (0:48-0:52) mm long, 0-19 (0:18-0:20) length of wing, with two main apical truncate
sensilla, adorals small. Cibarium with 14 teeth and about two or three rows of fore teeth. Pigment patch
rounded anteriorly. Pharynx with faint ridges. Hypopharynx (and mandibles) curled like grass, with very
low teeth and a smooth tip. Antenna 3 = 0-48 (0:45-0:52) mm long, 0-18 (0:17-0:20) length of wing,
1-23 (1:13-1:28) times length of 4+ 5, 0-19 (0-88-1-02) length of labrum, two ascoids on segments 3-15,
that on 4 delicate with long spur, no papilla on 5. Mandible pointed. Maxilla with eight broad lateral
teeth and 88 ventrals, and a dental depth of 0:29 mm, palpal ratios about 10: 25 : 31: 25:54. Scutum
and pleuron mainly pale. Wing length 2-64 (2:53-2:83) mm, 3-0 times width, R2/Re+3 1°83 (1:53-2:06),
R, overlap/R, 0-75 (0:72-0:78), rather constant. Abdominal tergites 5 and 6 with a few hind erect hairs.
Spermatheca oblong without collar.
$. Labrum 0-37 (0:36-0:37) mm long, 0-15 (0:14-0:15) length of wing. Cibarium with about ten
irregular hind teeth and about 40 scattered fore teeth. Pharynx with faint ridges. Antenna 3 = 0-61
(0:56-0:63) mm long, 0:25 (0:24-0:26) length of wing, 1:10 (0:98-1:17) length of 4+5, 1-65 (1:59-1:72)
length of labrum, one ascoid on segments 3-15, that on 4 with a narrow spur. Wing length 2:47 (2:35-2:55)
mm, 3:3 times width, R,/Re+3 1°81 (1-36-2-11), R; overlap/R, 0-73 (0:70-0:77). Aedeagus bluntly pointed.
Genital filaments short. Paramere beaked. Coxite with about 50 hairs in brush which merges gradually
into the dorsal thick hairs.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, 1973-74 (A. B. Knudsen, J. Jeffery & colleagues), 17 °,
48 3; ten of each measured.
DISTRIBUTION. West Malaysia: Gunong Besout Forest Reserve (as above, one with tip of proboscis
clogged); Ulu Gombak (Lewis, 1957 : 167).
Sergentomyia (Neophlebotomus) nankingensis (Ho, Tan & Wu) comb. n.
(Map 9)
Phlebotomus nankingensis Ho, Tan & Wu, 1954 : 427. Syntypes 4 9, 2 3, CHINA: Nanking area, Purple
Mountains, 1952 (depository unknown) [not examined].
Of the 14 cibarial hind teeth of the female, one on each side is separated from the rest. The
following features are among those given by the author.
2. Cibarium with 14 hind teeth, the outer two on each side spaced and apart from the rest; a row of 12
fore teeth present; pigment patch pointed in front and notched behind. Pharynx with groups of spicules.
282
Antenna stated to have one ascoid on 3-15. Palpal formula 1-2-3-4—5. Wing: R,/ R23 1:24, R, overlap/R.
0:52. Spermatheca with some distal striations.
$. Cibarium with eight teeth and a small faint triangular pigment patch. Antenna with one ascoid on
segments 3-15. Wing: R2/R23 1:1, R, overlap/R, 0-5. Coxite: persistent hairs not indicated. Style with
two of spines at 0-76. Position of seta not shown.
Sergentomyia (Neophlebotomus) perturbans (de Meijere)
(Figs 165-177, Map 10)
Phlebotomus perturbans de Meijere, 1909 : 201; Sinton, 1924d : 1015; 1928c : 316, 320; Patton & Hindle,
1926 : 405, 410 [type said to exist in Amsterdam]; 1928 : 533, 542; Edwards, 1928 : 64 [?]. Lectotype
9, JAVA, designated by Quate, 1967 : 42 (ZMA) [examined].
Phlebotomus sylvestris Sinton, 1924d : 1017 [conditional name justified by ICZN Article 17(8)]; 1928c:
320; 1931c : 1209; 193le: 110; 1932a: 62; 1933e : 420; Raynal, 19355 : 257; 1936a : 361 [synonymy].
Syntypes 2 3, INDIA (depository unknown) [not examined]. Syn. n.
[Phlebotomus perturbans de Meijere; Patton & Hindle, 1926:409; 1928:542. Misidentification in
view of later selection of lectotype. ]
Phlebotomus demeijerei Nitzulescu, 1930 : 540. Syntypes 2 9, JavA (depository unknown) [not examined].
[Synonymized (= sylvestris) by Theodor, 1948: 111.]
Phlebotomus (Prophlebotomus) sylvestris Sinton; Parrot, 1940: 311; 1946: 71, 73; Parrot & Clastrier,
1952 : 167.
Phlebotomus (Sergentomyia) whartoni Lewis, 1957 : 167; Quate & Fairchild, 1961 : 121; Lewis & Wharton,
1963 : 121. Holotype 2, West MALAYsIA (BMNH) [examined]. Syn. n.
Phlebotomus (Sergentomyia) sylvestris Sinton; Quate, 19625 : 265 [synonymy].
Phlebotomus (Sergentomyia) perturbans de Meijere; Quate, 1967 : 42.
Sergentomyia (Rondanomyia) whartoni (Lewis); Lewis, 1973a : 251.
The female differs from all other species in having about nine main cibarial teeth arising from a
well-marked refractive area of the ventral wall. The coxite of the male is long, narrow and slightly
curved.
9. Eye 0:61 length of head. Labrum 0-24 (0:23-0:25) mm long, 0-11 (0:11-0:11) length of wing, first two
apical sensilla with conspicuous sockets, adorals well developed. Cibarium with eight or nine distinct
pointed teeth merging into about ten spicules on each side, arising from thick refractive band, fore teeth
absent, pigment patch dark reddish brown anteriorly and grey posteriorly, with transverse and oblique
line, bearing anteriorly about eight longitudinal lines; distinct cibarial bulge present. Pharynx less than
twice as wide posteriorly as anteriorly, with faint ridges bearing minute spicules. Hypopharynx with low
teeth and a toothed tip. Antenna 3 = 0-36 (0:32-0:39) mm long, 0-16 (0:15-0:17) length of wing, 0-91
(0:85-0:95) length of 4+5, 1-48 (1:38-1:57) length of labrum, two ascoids on segments 3-15, that on 4
with vestigial spur, no papilla on 5. Mandibles nearly pointed. Maxilla with eight broad lateral teeth and
35 well-marked ventrals, and a dental depth of 0:15 mm, palpal ratio 10 : 22 : 30: 24: 56. Scutum brown
and pleura pale except near coxae. Wing length 2-22 (2:11-2:29) mm, 3-1 times width, R./R:,3 0-79
(0:62-0:88), R, overlap/R, 0-47 (0:20-0:67). Abdominal tergites 5 and 6 with a few erect hairs on hind
margins. Spermatheca with transverse wrinkles, no collar and a protruding knob.
$. Labrum 0-18 (0:14-0:20) mm long, 0-91 (0:08-0:10) length of wing. Cibarium with eight irregular
denticles of varied shape. Pharynx with faint ridges. Antenna 3 = 0-53 (0:50-0:57) mm long, 0-28
(0:25-0:31) length of wing, 0:78 (0:67-0:85) length of 4+ 5, 3-14 (2-48-3-78) length of labrum, one ascoid
on 3-15. Wing length 1-90 (1:74-2:03) mm, 3-6 times width, R,/R2:3 0°67 (0:37-0°85), R, overlap/R,
0-36 (0:18-0:44), costal basal node very small. Aedeagus slender and tapering to round point. Paramere
beaked. Coxite narrow and curved outward, with diffuse patch of about 40 narrow hairs, and many
thick long hairs pointing backward. Style narrow. Genital filaments about four times length of pump.
The above description is based on ten of each sex from West Malaysia, Gunong Besout Forest
Reserve (1974).
The antenna, very long in this species (Parrot, 1946), and long hairs on the coxite are remi-
niscent of P. frondifer and could indicate life in dark surroundings.
Measurements of four females from Java, namely the lectotype, paralectotype (lacking head) and two
283
172
-025
Figs 163-174 Sergentomyia species. 163, 164, S. malayae: (163) 9, spermathecae; (164) 3, termi-
nalia. 165-174, S. perturbans: (165) 2 (Gunong Besout area), labrocibarium; (166) labrum tip of
same; (167) 2 (Tanjong Rabok), cibarium; (168) 2, cibarium of lectotype; (169) 2 (Sukna), cib-
arium; (170, 171) 2 (Gunong Besout area), cibarial hind teeth crushed, and isolated pigment patch
of same fly; (172) 2, hypopharynx tip; (173) 2 (Sukna), spermatheca; (174) ¢ (Gunong Besout
area), head.
284
¥
‘
125
178
0)
S
Pe SCGre
-025
184
LS
JQ)
1G!
Figs 175-186 Sergentomyia species. 175-177, S. perturbans, 3, terminalia, and coxite in mesad and
lateral view. 178, 179, S. purii, 2: (178) mandible; (179) maxilla. 180-186, S. quatei: (180) 2, head
pharynx; (184) 9, maxilla; (185) 9, sperma-
(181) 9, labrocibarium; (182) 9, cibarium; (183)
thecae; (186) 3, cibarium.
285
wing. Antenna 3 = 0-35 (0:33-0:36) mm long, 0-15 (0:15-0:16) length of wing, 0-89 (0-86-0-93) length of
4+5, 1:67 (1:25-1:39) length of labrum. Palpal ratio about 10: 21:28:20:45. Wing length 2-18
(1:99-2:26) mm long, about 3-3 times width, Ry/R:+3 0-70 (0:59-0:88), R, overlap/R, 0-52 (0:36-0:70).
These and other features agree well with flies from West Malaysia.
Measurements of four females from Sukna, India (3.vii.1908), are as follows. Labrum 0:29 (0:27-0:31)
mm long, 0-13 (0:12-0:14) length of wing. Antenna 3 = 0-32 (0:29-0:35) mm long, 0-14 (0:13-0-15)
length of wing, 0-90 (0-83-0-96) length of 4+5, 1-11 (1-06-1-18) length of labrum. Palpal ratio about
10: 24:32:21:50. Wing length 2-27 (2:23-2:31) mm, 3-3 times width, R2/R2,3 0-71 (0:64-0:78), R,
overlap/R, 0:45 (0:38-0:48). These figures agree well with data from Java except that antenna 3/labrum
is lower, but this could well be due to infraspecific variation.
When S. perturbans was named it was probably thought that all sandflies bit man, but there is
no evidence that it does so.
DISCUSSION ON THE CIBARIUM OF THE FEMALE. Nitzulescu (1930) described the cibarial armature
of Javanese S. perturbans (=demeijerei) as a double row of teeth, nine in each, one behind the
other. Sinton (1932a) studied Indian specimens (=sy/vestris) and figured about 25 hind teeth on
a concave line, and an apparently refractive area (the above-mentioned dark ventral patch) in line
with the ten central ones. Raynal (19355) described, in North Vietnam sylvestris, 20 strong and
very pointed hind teeth and, at the anterior edge of the dark pigment patch, a row of ten or 11
contiguous rectangles, apparently anterior incisor teeth, which were more or less apparent at
certain angles during mounting. Raynal & Gaschen (1935b) stated that the hind lateral teeth
were sometimes difficult to see, and that the rectangles appeared pigmented, apparently owing to
the pigment patch on which they were reported to be placed. Parrot & Clastrier (1952), studying
specimens (=sylvestris) from Cambodia, recognized 13 dimorphic hind teeth, namely nine
median long ones, curved slightly inward, and on each side narrow ones which appeared to num-
ber two but were hard to see and could be more numerous. They observed the dark rectangles,
apparently chitinous thickenings, but could not confirm Raynal’s impression of incisor teeth.
Quate (1967) described, in Javanese perturbans, eight long teeth with bases apparently embedded
in a dark nail-shaped pigment patch, with no evident fore teeth, the armature being unlike that
of any other Asian species which he knew.
After studying cibaria from the three above-mentioned countries, mounted in Berlese’s medium
without pressure, and removing the pigment patch from a few specimens, I find that the differ-
ences in descriptions are due to several causes. Compression has sometimes increased the number
of observed hind teeth. The thick hind part of the inter-arcal area produces a refractive lens-like
effect which masks the structure. The lines on the pigment patch (Nitzulescu’s first row of ‘teeth’)
have been taken for teeth or bases of teeth; this can easily happen because in a cibarium at rest
the arched dorsal and ventral walls are almost in the same optical plane.
The lines on the pigment patch (also seen in S. reidi) appear to be imprints of the central
cibarial teeth. Among specimens examined in detail were three pinned ones from Sukna, India
(3.vii.1908, labelled P. perturbans, presented to the BMNH in 1927 by E. Brunetti, and recently
slide-mounted). They showed eight cibarial main teeth and, on each side, one vestigial tooth and
numerous denticles or spicules such as occur in several species of Sergentomyia and could well
be disregarded in descriptions. At the bases of the teeth is a dark area which appears to be a
thickening of the ventral wall of the cibarium. A hind bulge is conspicuous.
DISCUSSION ON SYNONYMY. S. perturbans was described at a time when some important taxonomic
structures and 98 per cent of the world’s sandfly taxa were still unknown. Species in distant lands
were wrongly identified as this species, and S. sylvestris was treated as a different species, but
now proves to be a synonym. For a time workers who examined specimens from de Meijere’s
collection tended to regard them as one species (they are here shown to comprise at least five),
and there was little attempt to distinguish the type-series from other specimens. Two specimens
of what proves to be the type-form were described as a new species, demeijerei. Confusion has
reigned for over 65 years, and has been referred to in part by Patton & Hindle (1928), Sinton
(1928c), Raynal & Gaschen (1936c), Perfil’ev (1968) and Lewis & Dyce (1976).
286
The situation is clarified, it is hoped, in the following notes on specimens which, except num-
bers 5, 6, 15 and 16, are in the ZMA. All were collected in Java in 1908, 1909 and 1910 by Edward
Jacobson (a private collector who lived in Djakarta, then Batavia, and Semarang) and are females
except number 6.
1. P. stantoni, Semarang, xi.1909.
2 and 3. P. stantoni, Semarang, ii.1910.
4. S. (Parrotomyia) mangana or ally, Semarang, ii.1910. The arc bearing the cibarial teeth is
nodular.
5. S. sp., probably balica, no locality or date recorded. Cibarium figured by Patton & Hindle
(1928 : 542) as co-type. This and number 6 were accepted as perturbans by Sinton (1928c : 320).
6. S. sp., possibly near balica, 3, no locality or date recorded. Terminalia figured by Patton
& Hindle (1928 : 544) as co-type, name in legend wrong. Aedeagus pointed, paramere hooked,
style with all spines terminal.
7. S. (Neophlebotomus) balica, Djakarta, xi.1908, labelled ‘Patton, vis.’ Wing length 1-97 mm.
8 and 9. S. balica, Semarang, i.1910. Wing length 1-97 and 1:98 mm.
10 and 11. S. balica, Semarang, ii.1910. Wing lengths 1-94 and 1-98 mm. Average length for
numbers 7-11 is 1-97 (1:97-1:98) mm.
12. S. (Neophlebotomus) Djakarta sp., Djakarta, xi.1908, labelled ‘Patton vis. . . . P perturbans
det. de Meijere’, evidently by Patton. Antenna 4-16 and abdomen missing. Wing length 1-68 mm.
13 and 14 (others presumably existed but were not mentioned by describer). S. perturbans,
Djakarta, November and Semarang, June, described by de Meijere (1909). His mention of colour
differences between Djakarta and Semarang specimens foreshadowed what we now know, that
he was dealing with at least three species. His measurements for wing length suggested that
perturbans was a small species and were used by Sinton (1924d : 1016) to differentiate provision-
ally an Indian sandfly, sylvestris, from it. He reinforced this view (1928c : 320) on the strength
of drawings of specimens 5 and 6 by Patton and Hindle. Despite de Meijere’s evidently careful
drawing of the wing, his lengths prove unreliable, even if adjusted to show the length seen in the
figure. This adjusted length is about 0-86 of the wing lengths as measured in the present paper.
No original type-labels now exist, and Quate’s remarks (1967, published several years after
inspection of the specimens) are not clear. The specimen chosen by him as lectotype, mounted
in Euparal, and here numbered 13, retains the label ‘Patton vis.’ and is presumed to be one of
two labelled by de Meijere as ‘type’. Apart from this, neither of specimens 13 and 14 retains an
original label. Quate’s paper merely gives the locality as Java and the month as probably Novem-
ber. His small-scale drawing of the cibarium (perhaps made before the specimen had cleared)
differs from the appearance of Fig. 169 which, with other characters, shows that the lectotype is
the same taxon as that known for many years as S. sylvestris. The wing length is 2-25 mm.
Specimen 14 (wing length 1-99 mm) was labelled as paratype by Quate but not mentioned in
his paper, and it is stated to come from Java and has no head. It is here treated as the para-
lectotype.
15 and 16. S. perturbans, Semarang, vii.1909. Depository, if any, unknown. These were treated
by Nitzulescu (1930) as being found among co-types and described as P. demeijerei.
for numbers 13, 14, 17 and 18 is 2-18 (1:99-2:26) mm.
The West Malaysian whartoni is recognized as a synonym after inspection of perturbans from
Java and India.
MATERIAL EXAMINED.
India: Sukna, 4 °. Java: Djakarta and Semarang, 4 °. West Malaysia: Gunong Besout Forest Reserve,
1974 (A. B. Knudsen & colleagues), 14 2, 24 3; Tanjong Rabok, x, xi.1969, 3 2; Ulu Langat Forest
Reserve, 25.vii.1968, 1 2, 2 ¢ (A. Rudnick).
287
DISTRIBUTION. Bangladesh: Sylhet (Sinton, 1932a : 71). Burma: Rangoon (Sinton, 1928c : 320; 19324: 71).
Cambodia: Phnom Penh (Parrot & Clastrier, 1952 : 153). India: Jorhat (Sinton, 1924d : 1027); Doloi
Valley, Sukna (Sinton, 1928c: 200). Java: Djakarta, Semarang (de Meijere, 1909 : 201; Nitzulescu,
1930 : 541). Laos: Vientiane (Quate, 19625 : 265). Vietnam (North): Bim Son, Cho Ganh, Dong Giao,
Le Mi (Raynal, 19355: 261); Bui Huy Tin, Phu Oc (Raynal, 1936a : 362). West Malaysia: Gunong
Besout area (as above); Lubok Paku (Lewis, 1957 : 167); Gua ’Che Yatim (Quate & Fairchild, 196i : 211);
Tanjong Rabok (x, xi.1969, 3 9), Ulu Langat Forest Reserve (25.vii.1968, 1 9, 2 3) (A. Rudnick).
S. perturbans (=sylvestris) was taken by Sinton in the Darjeeling district of India, where it
seemed to be associated with foothill jungle and Raynal (1936a : 361) noted that it occurred at
quite a low altitude in India and Indo-China.
Sergentomyia (Neophlebotomus) purii (Sinton)
(Figs 178, 179, Map 10)
Phlebotomus purii Sinton, 1931d: 1203; 193le:110; 1932a:60; 1933c: 421. LECTOTYPE 9, here
designated [examined]. INDIA: Sukna, viii.1928 U. M. Puri), labelled ‘type 9’,
Phlebotomus (Prophlebotomus) purii Sinton; Parrot, 1940 : 311; 1946: 72.
Sergentomyia purii (Sinton); Theodor, 1948 : 11.
Sergentomyia (Rondanomyia) purii (Sinton); Theodor, 1958 : 2 [maxilla]; Lewis, 1973a: 251.
The long narrow pigment patch of the female is rather like that of S. Jinearis but the curved row
of teeth and the spermatheca are quite different.
DISTRIBUTION. India: Marianbari, Sukna, Tindharia (Sinton, 1931d: 1203).
S. purii was first found in forest tree-holes at the base of the Himalaya in Darjeeling District
(Sinton, 1932a).
Sergentomyia (Neophlebotomus) quatei Lewis sp. n.
(Figs 180-188, Map 10)
The species is distinguished from all related ones by its cibarial armature, which slightly resembles
that of S. silvatica, but S. quatei differs in having no pigment patch and a greater antenna 3/
labrum figure. The armature of S. quatei is rather like that of the Australasian S. brachycornuta
(Fairchild) which has a pigment patch and a shorter antenna 3.
2. Eye 0-71 length of head. Head and most of labium dark. Labrum 0-21 (0:21-0:23) mm long, 0-10
(0:09-0:10) length of wing, apical sensilla normal, subapicals vestigial, adorals large and some of them
accompanied by posteriorly converging lines. Cibarium with two middle teeth set backward, and about
five small ones separate from them on each side, pigment patch absent, faint bulge present. Pharynx
unarmed. Hypopharynx with vestigial serrations. Antenna 3 = 0-49 (0-46-0:52)mm long, 0:22
(0:22-0:24) length of wing, 0-86 (0:81-0:90) length of 4+5, 2-30 (2:21-2:36) length of labrum, ascoids
delicate and difficult to see against brown antenna, two with short blunt spurs seen on segment 3, and one
each on 3-10 and 12, many rather similar colourless hairs present, ascoid on 4 about 0-24 length of seg-
ment, no papilla on 5. Maxilla with nine low lateral teeth, and 32 small ventrals, dental depth 0-12 mm,
palpal ratio about 10: 14:23:16:41. Scutum, lower part of pleura, and coxae dark. Wing length
2:15 (2:15-2:23) mm long, 3-2 times width, R./Re+3 1°30 (1:05-1:50), R, overlap/R, 0-63 (0:40-0:68).
Spermatheca with protruding knob and internal spicules which are just visible in freshly mounted speci-
mens.
$. Labrum 0-18 (0:17-0:19) mm long, 0-10 (0:09-0:11) length of wing. Cibarial teeth very variable,
usually one to four large ones in centre and about six to ten small ones on each side, pigment patch absent.
Pharynx unarmed. Antenna 3 very long, 0:58 (0:54-0:64) mm, 0-32 (0:28-0:35) length of wing, 0-78
(0-68-0-84) length of 4+5, 3-18 (2:88-3:48) length of labrum, one ascoid on segments 3-10 (others
not seen). Wing length 1-82 (1:71-2:01) mm, 4 times width, R,/R2+3 1:16 (0-91-1-58), R, overlap/R,
0-54 (0:45-0:64). Aedeagus tapering to blunt point. Paramere with very unequal hairs. Coxite with
isolated sub-basal patch of about 31 hairs.
288
*05
o o—o- o—o—_o— oo +
a
198
=
=
196
+025
Figs 187-200 Sergentomyia species. 187, 188, S. quatei, 3: (187) terminalia; (188) paramere from
above. 189, 190, S. Sepilok sp., ¢: (189) tip of labrum from below, with upper hairs on right;
(190) coxite and style. 191, S. silvatica, 2 (Thailand), spermatheca. 192-200, S. tambori, 2: (192)
labrocibarium; (193) tip of labrum; (194) subapical and first adoral sensilla; (195) left tenth adoral
sensillum; (196) cibarium; (197) pharynx; (198) maxilla; (199) wing; (200) spermathecae.
289
MATERIAL EXAMINED.
Holotype 2, Borneo (Sabah): Sepilok, 7.ii.1972 (D. J. Lewis) (BMNH).
Paratypes. Same data, 7-15.ii.1972, 7 9, 27 3; 8 2 and 10 3 measured. (All in BMNH.)
Sergentomyia (Neophlebotomus) sp. (Rabok)
(Map 10)
The cibarial fore teeth are about four deep in a broad band, and the aedeagus is slender.
$. Labrum 0:25 (0:24-0:26) mm long, 0-14 (0:13-0:15) length of wing. Cibarium with three pointed hind
teeth and a few denticles on each side, and three rows of fore teeth, each with about ten. Pharynx with
faint ridges. Antenna 3 = 0-41 (0-41-0-41) mm long, 0-23 (0:23-0:24) length of wing, 1-18 (1-17-1-19)
length of 4+5, 1-66 (1:59-1:72) length of labrum, one ascoid on segments 3-15, that on 4 about 0-63
length of segment, with short blunt spur. Scutum and pleura very pale. Wing length 1-75 (1-71-1-79) mm,
about 3-1 times width, R,/Rj,, 1:39 (1:37-1:42), R, overlap/R, 0-63 (0:59-0:66). Aedeagus with tip
slightly turned up. Coxite with diffuse patch of about 30 thin hairs; style with seta and two of spines at
about 0-6.
In the absence of females this species is left without a formal name.
MATERIAL EXAMINED.
West Malaysia: Tanjong Rabok, 27.xii.1968 (A. Rudnick), 2 2.
Sergentomyia (Neophlebotomus) sp. (Sepilok)
(Figs 189, 190, Map 10)
The coxite brush has over 52 hairs, more than in S. hitchensi, and the seta on the style is at 0.4.
$. Rather dark, antennae with contrasting pale joints. Labrum 0-18 (0:17-0:20) mm long, 0-08 (0-08-0-09)
length of wing, with stout upper brush-hairs. Cibarium with about six hind teeth and about six fore teeth,
each in arow. Antenna 3 = 0-53 (0:-48-0-60) mm long, 0-24 (0-23-0-26) length of wing, 1-22 (1-08-1-29)
length of 4+ 5, 2-93 (2:79-3:10) length of labrum; ascoid on 4 = 0-2 length of segment. Palpal segment
4 subequal to or slightly longer than 3. Wing length 2-21 (2:13-2:27) mm, 3-6 times width, R,/Rois
1:77 (1:52-1-89), R, overlap/R, 0-67 (0:54-0:79). Coxite with over 52 hairs in brush; style with seta at
0-4 and the two subterminal spines at about 0-85.
In the absence of a female the species is not formally named.
MATERIAL EXAMINED
Borneo (Sabah): Sepilok, 15, 16.ii.1972 (D. J. Lewis), 4 3.
Sergentomyia (Neophlebotomus) silvatica (Raynal & Gaschen)
(Fig. 131, Map 10)
Phlebotomus sylvaticus Raynal & Gaschen, 1935d: 592 [3 in fig. 4 not this sp. according to Quate,
1926b : 264]. Syntypes 3 2, 2 ¢, VIETNAM (NorTH) (depository unknown) [not examined].
Phlebotomus silvaticus Raynal & Gaschen; Anonymous, 1935 : 779 [justified emendation (ICZN Article
32(B))]; Raynal, 19355 : 265.
Phlebotomus (Prophlebotomus) silvaticus Raynal & Gaschen; Parrot, 1940: 312; 1946: 71; Parrot &
Clastrier, 1952 : 162 [a 3].
Sergentomyia silvatica (Raynal & Gaschen); Theodor, 1948 : 112.
Phlebotomus (Sergentomyia) silvaticus Raynal & Gaschen; Lewis, 1957 : 167 [a 3].
Phlebotomus (Sergentomyia) sylvaticus Raynal & Gaschen; Quate, 1962b : 262 [description and fig. of
antenna 3 different, pigment patch of 3 variable].
Sergentomyia (Rondanomyia) sylvatica (Raynal & Gaschen); Lewis, 1973a : 251.
The cibarium of the female has about four to six teeth in the centre and a patch of eight to ten
smaller ones on each side.
290
2 (from Thailand). Labrum 0:22 (0:21-0:22) mm long, 0-09 (0:09-0:09) length of wing, adoral sensilla
much as in S. hamidi and S. quatei. Cibarium with two main teeth and no visible pigment patch. Hypo-
pharynx with low undulations. Antenna 3 = 0-40 (0-39-0-41) mm long, 0-17 (0-16—-0-17) length of wing,
0-95 (0-89-1-02) length of 4+5, 1:86 (1:79-1:94) length of labrum. Maxilla with nine lateral and 29
ventral teeth, dental depth 0:10 mm. Wing length 2-38 (2:35-2:41) mm, R,/R2,; 0:73 (0:53-0:92), R,
overlap/R, 0-52 (0:47-0:57).
MATERIAL EXAMINED.
Thailand: Ban Bon Dan, 12.xii.1975 (D. J. Gould), 2 9.
CommMeENTs. The Thailand females appear to be this species despite certain differences from
Raynal’s and Quate’s descriptions. S. hamidi and S. quatei have a rather similar labrum and
cibarial armature but differ in the spermathecal and other features.
Quate disregards the authenticity of Raynal’s male because it has a cibarial armature like that
of the female. The matter should not be regarded as settled because the sexes of the related S.
quatei are very similar in this respect.
DISTRIBUTION. Cambodia: Phnom Penh (Parrot & Clastrier, 1952: 155). Laos: Vientiane (Quate,
19626 : 264). Thailand: Ban Bon Dan (as above). Vietnam (North): Dong Giao, Nao Phu, Phu Oc (Raynal,
19355 : 269). West Malaysia: Rantau Panjang (? Lewis, 1957 : 67).
Sergentomyia (Neophlebotomus) tambori Lewis & Jeffery sp. n.
(Figs 192-203, Map 10)
The cibarium of the female has about 11 hind teeth (fewer than in S. iyengari), one irregular
row of fore teeth, and a broad anterior projection on the pigment patch.
S. tambori, like S. iyengari, somewhat resembles the African S. decipiens and S. dureni but
has a broad spermatheca.
9. Labrum 0:17 (0:17-0:17) mm long, 0:10 (0:09-0:10) length of wing, with normal distal sensilla, sub-
apicals with anterior pair small, adorals large with pegs pointing mesally, and cibarials less large with
longer pegs. Cibarium with about 11 large pointed teeth, the two sublaterals on each side very wide,
about 11 small fore teeth present in an irregular row, pigment patch pale brown, about 0-7 width of
cibarium and having broad anterior projection with rounded tip. Pharynx with a few sparsely spiculate
lines. Hypopharynx with about eight scarcely visible undulations on each side. Antenna 3 = 0-42
(0:40-0:44) mm long, 0:24 (0:23-0:25) length of wing, 1:24 (1-21-1-26) length of 4+5, 2-46 (2-42-2-50)
length of short labrum, two ascoids on segments 3-15, that on 4 = 0-4 length of segment, no papilla
on 5. Mandibles pointed. Maxilla with four low lateral teeth, 16 distinct ventrals, and a dental depth of
0:06 mm, palp with segment 1 small, ratio 10 : 27: 53 : 56: 89. Scutum pale brown and pleura pale.
Wing length 1-77 mm, 3-2 times width, R, (very long)/R2+ 3 3-31 (3-05-3-55), R, overlap/R, 0-79 (0:76-
0-81). Spermatheca broad and oblong, striated, with small knob in shallow pit, and faint ducts.
3. Head missing. Scutum pale reddish brown and pleura mainly pale. Wing length 1:59 mm, 3-8 times
width, R./ R23 1-00, R, overlap/R, 0-71. Genital filament about 4-1 times length of pump which has large
barrel. Aedeagus nearly parallel sided, mainly dark, with round tip. Paramere with pointed tip which
scarcely turns downward. Coxite with no differentiated brush. Style: dorso-mesad spine thick; dorso-
lateral spine thin; ventro-mesad spine (the only non-terminal one) rather thin and somewhat mesad;
ventro-lateral spine thick; seta at 0-65.
MATERIAL EXAMINED.
Holotype ?, West Malaysia: Gunong Besout Forest Reserve, 3.v.1974 (K. A. Tambor) (BMNH).
Paratypes. Same data as holotype, 3.v.1974, 1 3; 5.ii.1974 (J. Jeffery & colleagues), 1 9. (Allin BMNH.)
Sergentomyia (Neophlebotomus) tonkinensis (Raynal & Gaschen)
(Map 10)
Phlebotomus tonkinensis Raynal & Gaschen, 1935h: 742 [figs in 1935g]; Raynal, 19355: 273 [figs].
Holotype 2, VIETNAM (NortTH) (depository unknown) [not examined].
Phlebotomus (Prophlebotomus) tonkinensis Raynal & Gaschen; Parrot, 1940 : 312; 1946: 71.
291
Sergentomyia tonkinensis (Raynal & Gaschen) Theodor, 1948 : 112.
Sergentomyia (Rondanomyia) tonkinensis (Raynal & Gaschen) Lewis, 1973a : 251.
This species has neither fore teeth nor pigment patch.
DISTRIBUTION. Vietnam (North) (rare): Phu Doan (Raynal, 19355 : 274; 1936a : 363).
Sergentomyia (Neophlebotomus) traubi (Lewis) comb. n.
(Map 10)
Phlebotomus traubi Lewis, 1957: 169; Quate & Fairchild, 1961: 216. Holotype 9, West MALAYSIA
(BMNH) [examined ].
The female has fold-like cibarial teeth unlike those of other species. The species is placed in
Neophlebotomus owing to the structure of the terminalia of the male.
g. Labrum 0:18 mm long, 0-11 length of wing. Cibarium with 12 fold-like teeth, longer in the centre, and
no pigment patch. Pharynx with a few faint ridges. Antenna 3 = 0-53 mm long, 0:33 length of wing,
2:94 length of labrum; ascoid on 4 = 0-15 length of segment; segment 5 and rest missing. Wing length
1-61 mm, R2/R2,3 0°75, R, overlap/R, 0-35. Genital filament 3-0 length of pump. Aedeagus nearly
parallel-sided, and tapering rather abruptly to a broadly rounded tip. Paramere slightly beaked. Coxite
long and narrow, with hairs little differentiated except for four big ones in a row near tip. Style narrow
with two spines at 0-67, a seta at 0-73 and two spines at tip.
MATERIAL EXAMINED.
West Malaysia: Bukit Ibam, 1.x.1961 (L. W. Quate), 1 3.
DISTRIBUTION. West Malaysia: Bukit Ibam (L. W. Quate via BPBM), commonest species, 41 92, 1 3;
Teranggan (Quate & Fairchild, 1961); Ulu Gombak (Lewis, 1957). Borneo (Sabah): Kalabakan River
(Quate & Fairchild, 1961).
Sergentomyia (Neophlebotomus) zeylanica (Annandale)
(Fig. 204, Map 10)
Phlebotomus zeylanicus Annandale, 1910a: 60; 19116: 203; Brunetti, 1912: 215; Sinton, 1924e : 1029
[sexes misassociated, ? valid species, described; 3 was P. argentipes]; 1928c : 319; 1931le: 110; 1932a:
61; 1933e : 420; Theodor, 1938a: 261 [2 3, ¢ differs from previous descriptions]; Mitra, 1953b : 162.
Lectotype ?, SRI LANKA (Zoological Survey of India), designated by Quate, 1962c : 158 [not examined].
Phlebotomus chalami Young & Chalam, 1927 : 849 [conditional name]. Syntypes 2 3, INDIA (sent to
former Central Research Institute, Kasauli) [not examined]. [Synonymized by Sinton, 1928c : 319.]
Phlebotomus (Prophlebotomus) zeylanicus Annandale; Parrot, 1946 : 70, 71.
Phlebotomus (Sergentomyia) zeylanicus Annandale; Quate, 1962c : 158, 160 [spermatheca].
Sergentomyia (Rondanomyia) zeylanica (Annandale); Lewis, 1973a : 251.
The female of S. zeylanica differs from S. malayae in having a shorter labrum.
2 (extra facts). Labrum 0-31 mm long, 0-15 length of wing (1:20 mm). Antenna 3 = 0-31 mm long,
0-16 length of wing, 1-14 length of 4+5, 1-01 length of labrum. Maxilla with three lateral and 30 ventral
teeth, dental depth 0-11 mm.
MATERIAL EXAMINED.
Sri Lanka: Peradeniya, 30.iv.1914, 1 9.
DISTRIBUTION. India: Kulathurpuzha (BMNH); Dehra Dun area, Naini Tal area (Rao et al., 1973);
Bombay, Gauhati (Sinton, 1928c:319); Marianbari (Sinton, 193le: 107); Darjeeling (Theodor,
1938 : 263). Sri Lanka: Peradeniya (Annandale, 1910a : 59, fig. 166, 25.viii.1910); Depanama, Kalgoda,
Katuwawala, Maharagama (Theodor, 1938 : 262).
S. zeylanica appeared to occur in many widely separated parts of India and to be a wild species
associated chiefly with jungle and moist warm conditions (Sinton, 1932a: 71).
292
Figs 201-214 Sergentomyia species. 201-203, S. tambori, 3: (201, 202) terminalia; (203) left style
in side-view. 204, S. zeylanica, 2 (Sri Lanka), spermatheca. 205, 206, S. bailyi, 2 (Andaman
Islands): (205) labrocibarium; (206) maxilla. 207-214, S. cheongi, 2: (207) labrocibarium; (208)
teeth; (209) teeth and pigment patch in postero-ventral view; (210) teeth and pigment patch
crushed; (211) teeth and pigment patch in thick section, anterior view; (212) crushed teeth;
(213) pharynx; (214) spermatheca.
293
The nicnic-group
Sergentomyia nicnic group Theodor, 1948 : 102.
The minute cibarial teeth are a feature of this group in which Theodor placed S. bailyi, S.
kachekensis, S. nicnic and one African species which may not belong to it.
Sergentomyia bailyi (Sinton)
(Figs 205, 206, Map 11)
Phlebotomus bailyi Sinton, 19316: 821; 1932a: 60; Parrot, 1946: 72. Syntypes 2 3d, INDIA (depository
unknown) [not examined ].
Phlebotomus bailyi var. campester Sinton, 19316 : 822; 1931d: 104; Raynal, 1935a: 369; 1935b : 277;
Raynal & Gaschen, 1934c : 563; 1934e : 858; Theodor, 1938 : 268; Causey, 1938 : 487; Yao & Wu,
1940 : 782; 19415: 77. [Synonymized by Quate, 19625 : 262.]
Phlebotomus (Prophlebotomus) bailyi var. campester Sinton; Parrot & Clastrier, 1952: 155 [including
abnormality J.
Phlebotomus smithi Mitra & Roy, 1952a : 187 [2]; Mitra, 1953a : 473 [3]. Holotype 9, INDIA (depository
unknown) [not examined]. Syn. n.
Sergentomyia (Sergentomyia) bailyi (Sinton); Theodor, 1948 : 112; Lewis, 1967 : 38.
Phlebotomus (Sergentomyia) bailyi Sinton; Quate, 19626 : 260, 261, 262, 264.
In the female the cibarial cornua are large, the pigment patch is small or absent, and the very
small teeth tend to be arranged in rows especially at the sides, the pharynx has spiculate ridges,
and the spermatheca (Sinton, 1932a : 60) narrows at its apex.
The female, and probably the male, of S. smithi appear to be S. bailyi, and apparent differences
to be caused by mounting methods and variation.
2 (extra facts). Mandibular main teeth 2-0 um wide. Maxillary ventral teeth large.
DIsTRIBUTION. Andaman Is.: South Andaman, Chinya Tapu, 12.ii.1970 (N. L. Kalra), 2 9. Cambodia:
Phnom Penh (Parrot & Clastrier, 1952 : 153). China: Aihsien, Kachek, Kan-en, Lingmen, Lingshui,
Linko, Mencheong, Nodoa, Paoting, Wanning (Yao & Wu, 1940 : 797; 19415 : 77). India: Ajmer (Jaswant
Singh, 1933); Badam Pahar, Itaunia, Patna, Vellore (BMNH); Aurangabad, Jalna, Patan (Farooq &
Qutubuddin, 1945 : 85); Bundi area, Jaipur, Kota area, Sirohi area (Kaul et al., 1973 : 532); Poona
(Mitra & Roy, 1952a, S. smithi); Hyderabad (Qutubuddin, 1944 : 208); Madras (Rathnaswamy & Rama
Krishna, 1954); Barhi, Bissem Cuttack, Chandigarh, Chhindwara, Dehra Dun, Hardwar, Hazaribagh,
Itarsi, Kamptee, Karnal, Kasauli, Khandwa, Laharpur, Nagpur, Pachmarhi, Parasia, Pipariya, Roorkee,
Saharanpur, Titilagarh (Sinton, 19315 : 821); Bhavnagar, Rajkot, Sanawar, Wadhwan (Sinton’s notes);
Baraga, Hosur, Jog-Sagar area, Kannur, Kumsi, Poona, Wai (H. Trapido). Laos: Vientiane (Quate,
19626 : 262). Pakistan: Dehra Ismail Khan, Jandola, Kohat-Hangu valley, Lahore, Larkana, Pano Aqil,
Peshawar, Rawalpindi, Shikarpur, Tando Muhammad Khan, Tank, Taxla (Lewis, 1967 : 39). Thailand:
Bangkok (Sinton, 1931d: 104); in train (Causey, 1938: 487, 488); Pechaburi (Quate, 19625 : 262).
Vietnam (North): Bim Son, Cho Ganh, Cua Rao, Dong Giao, Kep, Phu Qui, Vin Thui (Raynal,
19355 : 282). Vietnam (South): Duc Pho (Raynal, 19355 : 282).
In India Sinton (19315) found that S. bailyi was widespread up to 1830m, but much less
numerous than S. babu, and that it was relatively more numerous in the hills. In Indo-China
Raynal (1936a : 351, 357) found S. bailyi south of 20° north.
Sergentomyia displicata (Quate & Fairchild)
(Map 11)
Phlebotomus (Sergentomyia) displicatus Quate & Fairchild, 1961 : 212 [3]. Holotype 3, BoRNEo (BPBM)
[not examined ].
The cibarium of the male has about 15 hind teeth and a double row of 20 fore teeth, and the area
between them and the arch is dark. The male differs from that of S. nicnic in having no pigment
patch and a longer antenna 3.
294
DISTRIBUTION. Borneo (Sabah): Kalabakan River (Quate & Fairchild, 1961 : 214).
Sergentomyia kachekensis (Yao & Wu)
(Map 11)
Phlebotomus kachekensis Yao & Wu, 1940 : 790 [3]; 19415 : 78. Holotype 3, CHINA (depository unknown)
{not examined ].
Sergentomyia kachekensis (Yao & Wu); Theodor, 1948 : 113.
This species, described from one male, was placed in this group by Theodor. It is left here, in
the absence of more material, although the original drawing showing diamond-shaped teeth
suggests that it may belong elsewhere.
DisTRIBUTION. China: Hainan Island (Yao & Wu, 19415 : 77); Kachek (Yao & Wu, 1940 : 790).
Sergentomyia nicnic (Banks)
(Map 11)
Phlebotomus nicnic Banks, 1919a:163; Sinton, 1928c:317; 1930a:165; 1931b:824; Manalang,
1930a : 169; Theodor, 1938 : 268. Holotype 3, PHILIPPINES (destroyed, according to Quate & Rosario)
[not examined ].
Phlebotomus (Prophlebotomus) nicnic Banks; Parrot, 1940 : 311; 1946 : 72.
Phlebotomus (Sergentomyia) nicnic Banks; Quate & Fairchild, 1961 : 214; Quate & Rosario, 1962 : 794.
In this small species the female has a weak armature of many small teeth, big cornua, prominent
hypopharyngeal teeth, and spermathecae narrow proximally, and the paramere of the male has
an unusually large tip.
DISTRIBUTION. Nusa Tenggara: Kabaru, Pedang Bay, Sumbawa (Lewis & Dyce, 1976 : 212). Philippines:
Los Banos (Banks, 1919a : 167; Sinton, 1930a : 165); Nueva Viscaya (M. D. Delfinado & D. E. Hardy,
via BPBM); Novaliches (Manalang, 1930a : 169); Pili (Quate, 1965 : 28); Bay, Jose del Monte (Quate
& Rosario, 1962 : 796).
UNGROUPED
It is difficult to classify this miscellaneous assemblage, but it is convenient to divide the females,
as in the key, into those with thick cibarial teeth, those with certain outstanding features, and
those with a rather simple row of equal teeth. A few species could, perhaps, be offshoots of
Parrotomyia or Neophlebotomus which have lost their subgeneric characters.
Sergentomyia angustipennis (de Meijere) comb. n.
(Map 11)
Phlebotomus angustipennis de Meijere, 1909 : 202; Annandale, 1910b : 52; 191la: 62; Sinton, 1928c:
322; Nitzulescu, 1930 : 545. Holotype, sex not given, JAVA (depository unknown [not examined].
The short description of S. angustipennis, meaning short-winged, refers to a few external charac-
ters and is accompanied by a figure of the wing, the narrowness of which suggests that the species
belongs to the genus Sergentomyia. An adequate description could be prepared if S. angustipennis
could be recognized during a survey of the local species.
DIsTRIBUTION. Java: Semarang.
Sergentomyia anodontis (Quate & Fairchild)
(Map 11)
Phlebotomus (Sergentomyia) anodontis Quate & Fairchild, 1961 : 220; Lewis & Wharton, 1963 : 120.
Holotype 3, West MALaysIA (BPBM) [not examined].
295
In this small species with a brown scutum and pale pleuron the cibarium of the female has spine-
like projections from a fold in the membrane above the sclerotized part, and a medium projection
over which is an inverted V-shaped bar, and tergite 8 has a lateral patch of hairs. The spermatheca
is long and tubular with a thick knob and is not differentiated from its duct which joins a com-
mon duct. The style of the male is slender with a seta at 0-7, one spine at 0-75, one subterminal
and one terminal. S. anodontis is closely related to the Chinese S. koloshanensis (Yao & Wu).
In a female from Betis the hypopharynx has definite low teeth, and the maxilla has nine lateral
teeth of moderate size, 29 strong ventrals, and a dental depth of 0-09 mm.
DIsTRIBUTION. West Malaysia: Betis (Lewis, 1957 : 121); Batu Caves (Quate & Fairchild, 1961 : 220).
Sergentomyia cheongi Lewis & Jeffery sp. n.
(Figs 207-214, Map 11)
The female differs from most species in its pear-shaped cibarial teeth, and from S. Josarcus in
having a very short inter-arcal area, a smaller antenna 3/labrum figure, palpal segment 3 shorter
than 4, and Newstead’s sensilla concentrated.
2. Labrum 0-19 (0:18-0:20) mm long, 0-09 (0:10-0:11) length of wing, narrow, with very small subapical
sensilla, few adorals and small cibarials. Cibarium with inter-arcal thick posterior area supporting nine
large pear-shaped teeth with long dorsal points which are invisible from below; pigment patch brown
with about eight longitudinal lines. Pharynx with narrow unarmed hind end. Hypopharynx with low
rounded teeth. Antenna 3 = 0-31 (0:30-0:34) mm long, 0:16 (0:16-0:17) length of wing, 1-27 (1-23-1-33)
length of 44+.5, 1-60 (1:50-1:72) length of labrum, two ascoids on segments 3-15, that on 4 about 0-46
length of segment, no papilla on 5. Mandible with wide-angled tip. Maxilla with 12 broad lateral teeth and
25 ventrals, dental depth 0:07 mm; palpal ratio about 10 : 21 : 33 : 48 : 98; sensilla concentrated on basal
quarter of 3. Scutum, pleuron and much of body reddish brown. Wing length 1-90 (1:79-1:99) mm,
3-5 times width, Ry/Ro+3 1:34 (0:84-2:09; minimum exceptionally low), R, overlap/R, 0-60 (0:52-0:71).
Spermatheca oblong with delicate duct.
The harrow-like cibarial armature is seen in this species, S. perturbans, and a few others.
MATERIAL EXAMINED.
Holotype 2, West Malaysia: Gunong Besout Forest Reserve, 4.ii.1974 (J. Jeffery) (BMNH).
Paratypes. Same data, 9 2 (BMNH). Ten 2 measured.
Non-paratypic material. West Malaysia: Bukit Ibam (L. W. Quate via BPBM), 1 °.
Sergentomyia dapsilidentes (Quate)
(Map 11)
Phlebotomus (Sergentomyia) dapsilidentes Quate, 1965 : 26. Holotype 9°, PHILIPPINES (BPBM) [not ex-
amined ].
In the female of this pale species there are 30 cibarial hind teeth in a compact row and about
80 fore teeth in five or six rows, and an unusually long palpal segment 4.
DisTRIBUTION. Philippines: Manucan, Zamboanga del Norte (Quate, 1965 : 28).
Sergentomyia delfinadoae (Quate)
(Map 11)
Phlebotomus (Sergentomyia) delfinadoae Quate, 1965:30. Holotype 9, PHILIPPINES (BPBM) [not
examined ].
The female of this large species has a brownish scutum and about 10-14 cibarial teeth like
barbed hooks, a rectangular pigment patch and a long antenna 3. The male has six to ten barbed
teeth. The species somewhat resembles S. exastis.
296
2 (extra facts). Labrum with the foremost adoral sensilla large, in three diagonal pairs. Hypopharynx
smooth. Maxilla with eight moderately broad lateral teeth and 17 distinct ventrals, dental depth 0-10 mm.
DIsTRIBUTION. Philippines: Cuernos de Negros (Quate, 1965 : 30).
Sergentomyia dentacea (Quate)
(Map 11)
Phlebotomus (Sergentomyia) dentaceus Quate, 1965 : 26. Holotype ?, PHILIPPINES (BPBM) [not examined].
In the female of this large species the cibarial teeth are very long and the pharyngeal teeth are
markedly posterior.
DISTRIBUTION. Philippines: Los Arcos (Quate, 1965 : 26).
Sergentomyia exastis (Quate)
(Map 11)
Phlebotomus (Sergentomyia) exastis Quate, 1965 : 33. Holotype 2, PHILIPPINES (BPBM) [not examined].
This is a large species and the female has no pigment patch and 14 inwardly sloping cibarial
teeth rather like those of the related S. delfinadoae and the African S. schwetzi.
DISTRIBUTION. Philippines: San Francisco (Quate, 1965 : 33).
Sergentomyia fanglianensis (Leng)
(Map 11)
Phlebotomus fanglianensis Leng, 1964 : 118, 127. Syntypes 2 3, CHINA (Lianming University, Shenyang)
[not examined ].
In the female the cibarium has no pigment patch and many small triangular teeth tending to
form three rows medially, the pharynx has finely spiculate ridges, and the smooth carrot-shaped
spermatheca has a deep pit. In the male the cibarial teeth tend to form three rows, Ro/Ro,3 is
over 1-5 (1-63-1-80), and the style has a seta, two subapical spines on a tubercle and two apical
spines.
DISTRIBUTION. China: Fanglian (Leng, 1964 : 127).
Sergentomyia hassani Lewis sp. n.
(Figs 215-221, Map 11)
The female is very like that of S. hitchensi but has a long row of palpal sensilla.
9. Eye 0-62 length of head. Labrum 0-19 (0:18-0:19) mm long, 0-08 (0:07-0:08) length of wing, with
adoral sensilla well developed. Cibarium with narrow hind end and deep notch (not seen in macerated
flies) in dorsal wall, inter-arcal walls dark, and a nearly straight row of small separate teeth, which are
blunt in ventral view and merge into spicules on cibario-pharyngeal membrane, and no pigment patch.
Pharynx unarmed and rather soft with variable shape. Hypopharynx completely smooth. Antenna
3 = 0-39 (0:38-0:40) mm long, 0:16 (0:15-0:17) length of wing, 1-35 (1-31-1-36) length of 4+5, 2-13
(2:09-2:16) length of labrum, two ascoids on segments 3-15, that on 4 being slender and about 0-43
length of segment, no papilla on 5. Maxilla with nine narrow lateral teeth, 25 ventrals, and a dental depth
of 0-09 mm; palpal ratio 10 : 12 : 26 : 27 : 63; sensilla scattered along most of length of 3. Scutum pale
reddish brown, pleuron mainly pale. Wing 2-42 (2:32-2:53) mm long, 3-3 times width, R2/R2+3 2°38
(2:19-2:56), R, overlap/R, 0:75 (0:74-0:76). Leg ratios: fore (0-96 mm), 50: 80: 46 ; hind (0-84 mm),
50 : 99 : 52; hind (0-94 mm), 50 : 104 : 71. Spermatheca subpyriform with delicate duct.
297
© ° 7) 226
a :
@D Q) ” Pep
g 0”
Y, MG
PO @o@2
sto
Figs 215-229 Sergentomyia species. 215-221, S. hassani: (215) 2, labrocibarium; (216) 9, cibarium;
(217) 9, tip of hypopharynx; (218) 9, palpal segment 3; (219) 9, spermatheca; (220) 3, cibarium;
(221) 3g, terminalia. 222-227, S. jamesi, 2: (222) labrocibarium; (223) cibarium; (224) pharynx;
(225) wing; (226) spermatheca; (227) pattern on contained egg. 228, 229, S. knudseni, 9: (228)
labrocibarium; (229) cibarium.
298
$. Labrum 0-17 (0-15-0-19) mm long, 0-08 (0-07-0-09) length of wing. Cibarium with deep hind notch
in soft tissue, about seven minute teeth, and no pigment patch. Pharynx unarmed. Antenna 3 = 0-46
(0:42-0:49) mm long, 0-22 (0:21-0:23) length of wing, 1-33 (1-28-1-38) length of 4+5, 2-78 (2:50-3-01)
length of labrum, one ascoid on segment 3-15, that on 4 being 0-27 length of segment. Wing 2-12 (2-01-
2:26) mm long, 3:5 times width, R2/R2,3 1:53 (1:07-1:88), R, overlap/R, 0:69 (0:65-0:77). Aedeagus
with narrow rounded tip, filament 3-3 times length of pump. Coxite with about 16 hairs in brush.
Style with long stout seta at 0.25, one spine at 0-65, one at 0-75 and two at tip.
ComMENTS. S. hassani, like many sandflies, has a mixture of apparently plesiomorphic and apo-
morphic features. The former comprise a large delicate pale body, well-developed labral adoral
sensilla, a cibarium with narrow hind end, small rather irregular teeth, and no pigment patch,
extra ascoid-like structures on the antenna of the male, narrow maxillary lateral teeth, scattered
Newstead’s sensilla, a broad wing with long R, and a simple spermatheca.
This species is named after Mr Abu Hassan bin Omar of Kuala Lumpur who collected many
sandflies.
MATERIAL EXAMINED.
Holotype ?, West Malaysia: Ulu Langat Forest Reserve, 3.v.1966 (A. Rudnick) (BMNH).
Paratypes. Same data, 2 9, 8 ¢ (BMNH).
Non-paratypic material. West Malaysia: Kuala Tahan (D. E. Hardy & T. C. Maa, via BPBM); Bukit
Ibam and Kuala Rompin area (L. W. Quate, via BPBM).
Sergentomyia heiseri (Manalang)
(Map 11)
Phlebotomus heiseri Manalang, 1930e : 299; Tonnoir, 1935 : 142; Raynal, 1935a: pls 10, 19; Quate &
Rosario, 1962 : 796. Lectotype 9, PHILIPPINES (on loan to BPBM), designated by Quate & Rosario
[not examined ].
The cibarium of the female has 12-18 long diamond-shaped hind teeth. It bears some resemblance
to species of Parrotomyia but has different teeth, and shape and venation of the wing.
DIsTRIBUTION. Philippines: Bigti, La Mesa, Novaliches, Tungkong Manga (Manalang, 1930e); Los Bafios
(Quate & Rosario, 1962 : 797).
Sergentomyia hitchensi (Manalang)
(Map 12)
Phlebotomus hitchensi Manalang, 1930d : 291; Tonnoir, 1935 : 142. Lectotype 2, PHILIPPINES (on loan to
BPBM), designated by Quate & Rosario, 1962 : 791 [not examined].
Phlebotomus (Sergentomyia) hitchensi Manalang; Quate & Rosario, 1962 : 791; Quate, 1965 : 30.
Sergentomyia (Rondanomyia) hitchensi (Manalang); Lewis, 1973 : 250.
The female has 12-15 triangular teeth in a regular row and no pigment patch. The species was
placed in the zeylanica group by Theodor (1948) but is now removed owing to its uncharac-
teristic cibarial teeth and close affinity to S. Josarcus.
DIsTRIBUTION. Philippines: Tungkong Manga (Manalang, 1930d : 291); Eran area (Quate, 1965 : 30).
Sergentomyia imitor (Quate)
(Map 12)
Phlebotomus (Sergentomyia) imitor Quate, 1965 : 35. Holotype 9, PHILIPPINES (BPBM) [not examined].
In this small pale species the cibarium of the female has 10-12 sharp spike-like teeth (a conver-
gent resemblance to S. clydei) and 24-40 main fore teeth in two or three rows. The small well-
defined coxite-brush of the male is at 0-3 and the style has two apical and two postmedial spines
and a seta at 0-4-0-6. The terminalia are like those of S. displicata but the head is different.
DISTRIBUTION. Philippines: Los Arcos (Quate, 1965 : 37).
299
Sergentomyia jamesi Lewis sp. n.
(Figs 222-227, Map 12)
The female differs from all other Sergentomyia in having no cibarial teeth except a few lateral
denticles.
2. Labrum 0-16 mm long, 0-11 length of wing, with several posterior adoral sensilla and large lateral
labro-cibarial sensilla. Cibarium with no teeth but a small group of denticles at each side; pigment patch
absent and arch faint. Pharynx with faint ridges. Hypopharynx smooth. Antenna 3 = 0-12 mm long,
0-08 length of wing, 0-91 length of 4+5, 0-79 length of labrum, two ascoids on segments 3-15, that on
4 = 0°55 length of segment, no papilla on 5. Maxilla with 11 strong lateral teeth, 28 ventrals and a dental
depth of 0:07; palp ratio 10: 21:29:35: 68; sensilla in compact bunch around 0:37. Scutum and
pleuron brown, inter-precoxal lobes narrow, mesanepisternum without hairs. Wing length 1-48 mm,
4-1 times width, R./R2,3 0°89, R, overlap/0-39. Leg ratios: fore (0-49 mm), 10 : 8-8 : 4-8; mid (0-54 mm),
10 : 10-5 : 5-5; hind (0-59 mm), 10: 12 : 6-3. Spermatheca oblong.
Egg sculpture (seen in the single gravid 9). Polygonal (as in some American species, Ward & Ready,
1975 : 128).
The species is named after Mr Samuel S. James, of the Arbovirus Research Laboratory field
staff, West Malaysia.
MATERIAL EXAMINED.
Holotype 2, Thailand: ‘Trang Prov., Khaophappha Khaoc Hang, 200-300 m, 3.1.1944’ (G. A. Samuelson,
via BPBM). Mapped in Trang area.
Sergentomyia kelantani (Lewis & Wharton)
(Map 12)
Phlebotomus (Sergentomyia) kelantani Lewis & Wharton, 1963: 123. Holotype 9, West MALAYSIA
(BMNH) [examined].
The female has nine large delicate cibarial teeth, with blackish pigment near their bases, antenna
3 = 0:19 mm long, and the spermatheca is a rigid thin-walled capsule bent double with a wide
duct. The style of the male has two terminal and two subterminal spines.
DISTRIBUTION. West Malaysia: Betis (Lewis & Wharton, 1963 : 123).
Sergentomyia knudseni Lewis & Jeffery sp. n.
(Figs 228-237, Map 12)
A pale species with, in the female, regular cibarial teeth on a convex row, few or no fore teeth,
a large bulge, maxilla without lateral teeth, and a broad wing with (in both sexes) short R, and
R,-overlap.
The females of several species have rather similar cibarial hind teeth. S. bukidnonis has fore
teeth and more hind ones, and S. franciscana has fore teeth and a shorter labrum. The three
Australian species, S. cidaria (Quate & Quate), S. crypta (Quate & Quate) and S. vanella (Quate
& Quate) each have at least one distinct row of fore teeth.
9. Labrum 0-19 (0:19-0:20) mm long, 0-12 (0:12-0:13) length of wing, with few adoral sensilla. Cibarium
with arch far back, pigment patch broad but faint, and a wide hind bulge; 24-27 uniform pointed teeth
present on slightly convex arch, and occasionally two fore teeth near each margin. Pharynx with marked
brown subterminal bulge, narrow hind end and no teeth. Hypopharynx with low undulations. Antenna
3 = 0-23 (0:21-0:25) mm long, 0-14 (0:13-0:16) length of wing, 1-09 (1:04-1:17) length of 4+5, 1:17
(1-05-1-26) length of labrum, two ascoids on segments 3-15, that on 4 being slender, difficult to see, 0-86
length of segment and reaching the next one, no papilla on 5. Mandible pointed. Maxilla without lateral
teeth but five vestigial specks, and 22 small ventrals, dental depth 0-07 mm; palpal ratio 10: 18 : 31: 35:
64; sensilla based on basal third of 3. Scutum brownish, pleuron mainly pale. Wing length 1-59 (1-50-1-66)
300
mm, three times width, R,/R..3 0°71 (0:54-0:90), R, overlap/R, 0-36 (0:27-0:41). Spermatheca oblong
and thin-walled with delicate duct joining short common duct.
$. Labrum 0:16 mm long, 0:11 length of wing. Cibarium with about eight distinct separate pointed
teeth on slightly convex arc, a strong bulge, and a pigment patch about two-thirds width of cibarium.
Pharynx almost unarmed and very narrow posteriorly. Antenna 3 = 0-31 mm long, 0-21 length of wing,
1-22 length of 4+ 5, 1-89 length of labrum, one ascoid on segments 3-15 (last two missing), that on 4 being
0-75 length of segment and reaching its tip. Wing length 1-45 mm, 3-6 times width, R,/Re,3 0°52, Ry
overlap/R, 0:15. Aedeagus with rounded tip. Coxite with nine narrow mesad hairs.
MATERIAL EXAMINED.
Holotype ?, West Malaysia: Gunong Besout Forest Reserve, 24.11.1974 (J. Jeffery) (BMNH).
Paratypes. Same data, 5.v-15.vi.1974, 13 2, 1 $; same data, 10.viii.1973 (R. B. Tesh), 1 9; Ulu Gombak,
20.viii.1960 (Abu Hassan bin Omar), 1 2. (Allin BMNH.) Ten 2 and 1 3 measured.
Sergentomyia lagunensis (Quate)
(Map 12)
Phlebotomus (Sergentomyia) lagunensis Quate, 1965 : 26. Holotype 2, PHILIPPINES (BPBM) [not examined].
A large species with most of the pleuron brown and, in the female, 26 spike-like cibarial teeth
and indistinct fore teeth.
DISTRIBUTION. Philippines: Los Bafios (Quate, 1965 : 26).
Sergentomyia losarcus (Quate)
(Map 12)
Phlebotomus (Sergentomyia) losarcus Quate, 1965 : 30. Holotype 2, PHILIPPINES (BPBM) [not examined].
In the female of this large dark species the slender cibarium has 8-10 stout teeth in a slightly
convex row and no fore teeth or pigment patch, and a dark area between the teeth and the
strong arch; the pharynx is unarmed and antenna 3 very long, and the spermatheca is sub-
pyriform. On the style of the male is a seta at 0-4 or 0-5, two spines at 0-8, and two distal. The
name is being treated as a noun.
The species is close to S. hitchensi, separable by the smaller number of differently shaped teeth
and by distribution, and they may be forms of one species according to Quate.
DISTRIBUTION. Philippines: Los Arcos, San Francisco area (Quate, 1965 : 30).
Sergentomyia maai (Quate & Fairchild)
(Map 12)
Phlebotomus (Sergentomyia) maai Quate & Fairchild, 1961 : 218. Holotype 2, BoRNEOo (BPBM) [not
examined ].
The cibarium of the female has 25 fish-hook-shaped teeth in an even comb and 14 fore teeth of
which the centre ones are the larger, the pharynx is unarmed, the ascoids very short, and the
spermatheca smooth and ovoid with a small head. On the style of the male two of the setae are
subterminal.
DIsTRIBUTION. Borneo: Tawau (Quate & Fairchild, 1961 : 218).
Sergentomyia mahadevani sp. n.
(Figs 238-242, Map 12)
In the female the cibarium has 50 long parallel teeth and a pigment patch with a broad fore end,
the hind end of the pharynx is broad and indented, with numerous small teeth, antenna 3 and
301
Figs 230-242 Sergentomyia species. 230-237, S. knudseni: (230) 9, sagittal section of fascicle and
cibarium; (231) 9, same of cibarium; (232) 2, same of an American species (Lutzomyia panamensis)
for comparison; (233) 2, pharynx; (234) 2, maxilla; (235, 236) 2, spermathecae; (237) 3, terminalia.
238-242, S. mahadevani, 2: (238) cibarium; (239, 240) pharynx; (241) wing; (242) spermathecae.
302
R, are long, and the spermathecae are nearly spherical. The nature of the cibarium and pharynx,
alone, suffice to distinguish this species from others.
9. Labrum 0-20 mm long, 0-09 length of wing. Cibarium with about 50 long parallel teeth and a pigment
patch with a broad fore end, hind bulge present. Pharynx with broad indented hind end and numerous
closely packed small teeth. Hypopharynx with low rounded teeth. Antenna 3 = 0-44 mm long, 0-20
length of wing, 1:27 length of 4+ 5, 2:16 length of labrum, two ascoids on antenna 3-15, that on 4 being
0-4 length of segment amd having a vestigial spur 7:5 um long, no papilla on 5. Maxilla with six not very
wide lateral and 18 ventral teeth and a dental depth of 0:06 mm, palpal ratio 10: 23 : 38:22: 113,
sensilla grouped near base. Scutum pale brown and pleuron pale. Wing length 2:24 mm, 3-5 times width,
R2/Ro+3 3°64, R; thick, R; overlap/R, 0-74. Spermathecae nearly spherical with delicate ducts.
MATERIAL EXAMINED.
Holotype 9, Thailand: Ban Bon Dan, 11.xii.1976 (D. J. Gould), light-trap with CO, in tropical evergreen
and deciduous forest (BMNH).
Sergentomyia montana (Sinton)
(Map 12)
Phlebotomus minutus var. montanus Sinton, 1924a : 809, 812; 1927c:949; 1927d: 26. Syntypes 9? 2,
INDIA and PAKISTAN (depository unknown except 1 9 in Indian Museum) [not examined ].
Phlebotomus montanus (Sinton); Sinton, 1927d:27 [cibarium]; 1927e:31; 1928c: 316; 19295: 174;
1932a : 61; 1933e : 422.
Sergentomyia montana (Sinton); Lewis, 1967 : 39; Artemiev, 19765 : 426.
The cibarium of the female has small lateral teeth in several rows, a pigment patch with broad
forward process and pointed tail, a narrow pharynx with many teeth, and a spermatheca narrow-
ing distally. The cibarium of the male has several rows of lateral teeth and a distinct narrow
pigment patch.
Artemiev proposed to include S. montana in Parrotomyia, and to modify the diagnosis of the
subgenus slightly.
DISTRIBUTION. India: Bhowali (BMNH); Chamba area, Tehri area (Rao et al., 1973); Kasauli, Naini Tal
(Sinton, 1924a : 812). Nepal: Syabrudens (L. W. Quate, 2.xi.1965). Pakistan: Bahrein, Gilgit, Khaira Gali,
Murree, Parkuta, Rawalpindi, Said Pur, Taxla (Lewis, 1967 : 40).
Sinton (1932a) found S. montana in the western Himalayan foothills at about 1830 m where
(1928c) it was common and replaced the lowland S. babu.
Sergentomyia morini (Raynal & Gaschen)
(Map 12)
Phlebotomus morini Raynal & Gaschen, 1935 f: 731 [3]; Raynal, 19355: 301; 1936a: 367. Syntypes
2 3, VIETNAM (SouTH) (depository unknown) [not examined].
Phlebotomus (Prophlebotomus) morini Raynal & Gaschen; Parrot, 1940 : 312; 1946 : 70.
The cibarium of the male has six or seven groups of small teeth, a few fore teeth, and no pigment
patch. R,/R,,; is 0-73-0-76, and two spines on the style are subapical. The genital filaments
are very long, 6:5—6-7 times as long as the pump.
DISTRIBUTION. Vietnam (South): Duc Pho (Raynal, 19355 : 303; 1936a : 367).
Sergentomyia neras (Quate)
(Map 12)
Phlebotomus (Sergentomyia) neras Quate, 1965 : 28. Holotype 2, PHILIPPINES (BPBM) [not examined].
This small species has antenna 3 very short and most of the scutum brown. The female has 14
303
inwardly sloping cibarial teeth in a compact row and two rows of 16 fore teeth, and a sub-
pyriform spermatheca.
DISTRIBUTION. Philippines: Eran area (Quate, 1965 : 18).
Sergentomyia sp. (Okinawa)
(Map 12)
Sergentomyia sp. Lien, 1975 : 298.
In the one female known the cibarium has 70 teeth, the pharynx has about ten short spicules,
R,/R»,3 is 48 and the spermatheca is subovoid.
DISTRIBUTION. Japan: Ryukyu Retto, Okinawa.
Sergentomyia pachystoma (Quate & Fairchild)
(Figs 243-250, Map 12)
Phlebotomus (Sergentomyia) pachystoma Quate & Fairchild, 1961 : 215. Holotype 9, BoRNEo (BPBM)
[not examined ].
This pale species was named after a thick proboscis. In females from Borneo the cibarium has
22 teeth and 18 fore teeth, the inter-arcal area is sclerotized and wrinkled, the ascoids have a
small spur and there appears to be only one ascoid on segments 11-15, and the spermatheca
narrows before the tip and has a deep pit. The male is described here for the first time from West
Malaysian specimens regarded as this species.
2. Eye 0-44 length of head. Labrum 0-24 (0:23-0:26) mm long, 0-14 (0-13-0-15) length of wing. Cibarium
with 24-2 (20-29) teeth and 19-0 (16-22) fore teeth, and a marked dorsal hind bulge, inter-arcal area
with a wide brownish area merging into the brown arch but scarcely a trace of oblique lines, pigment
patch rather broad and bearing imprints of fore and hind teeth. Pharynx at widest point about 0.81 width
of cibarium. Hypopharynx with rounded teeth less than half as high as wide. Antenna 3 = 0-29 (0-27-0-30)
mm long, 0-16 (0:15-0-18) length of wing, 0-13 (0:12-0:14) length of 4+5, 1-19 (1:07-1:25) length of
labrum, ascoids with small spur, two ascoids on segments 3-15, distal ones delicate and sometimes
difficult to see. Each mandible pointed, with wide teeth. Maxilla slender with small teeth, seven laterals
and 36 ventrals, and a dental depth of 0-12 mm. Wing length 1-77 (1:70-1:88) mm, about 3-1 times
length, R./R..3 1°52 (1:04-2:01), R, overlap/R, 0-60 (0:52-0:71). Leg ratios, for comparison with original
description; fore, 6: 5-4; mid, 6: 6-5: 3-4. Spermatheca narrowing before tip, with deep pit and long
knob.
g. Labrum 0-25 (0:24-0:26) mm long, 0-14 (0:14-0:14) length of wing. Cibarium with about six long
teeth and about 18 irregular fore teeth and no pigment patch. Pharynx with ridges and minute spicules.
Antenna 3 = 0:42 (0:40-0:46) mm long, 0-24 (0:22-0:25) length of wing, 1-18 (1:16-1:25) length of
4+5, 1-69 (1:62-1:77) length of labrum, one ascoid with short spur on segments 3-15, that on 4 slender
and about 0-6 length of segment. Wing length 1-79 (1-73-1-85) mm, 3-2 times width, R,/R,.3 1°33
(1:11-1°59), R, overlap/R, 0-57 (0:50-0:65). Aedeagus tapering with rounded tip. Paramere beaked.
Coxite with about 27 hairs in brush. Style with seta at 0-46, one spine at 0-42, one at 0-53 and two terminal.
ComMENTS. West Malaysian females differ from the Borneo description in several respects. In
the former the cibarial teeth are more numerous on average, inter-arcal ridges are inapparent,
the pigment patch (dissected out) proves to be a different shape, the pharynx is wider, the pro-
boscis does not appear thick and includes a rather long labrum, and ascoid distribution (prob-
ably), wing length and leg ratios are different. Differences, however, could be due to infra-
subspecific variation or to differences in mounting, and the two forms are treated as one species
till more is known of the Borneo form.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, 7.x.1973-15.vi.1974 (J. Jeffery & K. A. Tambor), 3 9;
Tanjong Rabok, 12—28.xi.1969 (A. Rudnick), 7 2, 10 3.
304
DISTRIBUTION. Borneo (Sabah): Kalabakan River, Ranau (Quate & Fairchild, 1961 : 216). West Malaysia:
Gunong Besout Forest Reserve, Tanjong Rabok (as above); Bukit Ibam (L. W. Quate, via BPBM).
Sergentomyia pooi (Yao & Wu)
(Map 12)
Phlebotomus pooi Yao & Wu, 1941a:71; 1941b:78 [3]. Syntypes 4 3, CHINA (depository unknown)
[not examined ].
The cibarium of the male has no pigment patch, 23-29 hind teeth and 10-15 fore teeth. Its wing
is about four times as long as wide, and R,/R,,3 averages 0-76. Satisfactory identification will
depend on finding the female.
DISTRIBUTION. China: Tienpao (Yao & Wu, 1941a: 71).
Sergentomyia reidi (Lewis)
(Figs 251-263, Map 13)
Phlebotomus (Sergentomyia) reidi Lewis, 1957: 169 [2]; Quate & Fairchild, 1961 : 220. Holotype 9,
West MALAysIA (BMNH) [examined ].
Behind the main cibarial teeth of the female is a remarkable group of teeth. Occasionally it may
be hidden, if a head is tilted, so that the cibarium looks rather like that of S. cheongi which,
however, has larger main teeth.
2 (extra facts). Labrum about 0-19 mm long and 0-10 length of wing, with well-developed adoral sensilla.
Cibarial main teeth arising from tough sclerotized band in front of hind margin of ventral plate; each
tooth pear-like in ventral view but supporting long upward-pointing spine which is continuous with dorsal
basal ridge of tooth; pigment patch with about 13 longitudinal lines. Hypopharynx with low blunt teeth.
Mandible with rounded tip and long low teeth. Maxilla with ten broad lateral teeth, 32 ventrals, and a
dental depth of 0-08 mm.
3 (first description). Labrum 0-16 (0:16-0:17) mm, 0-11 (0-11-0-11) length of wing. Cibarium with seven
or eight, slightly diverging and narrow, pointed hind teeth merging on each side into spicules of same
shape; behind main teeth a patch of 8-16 narrow bent pointed teeth; all teeth under low power appearing
spatulate owing to bending; pigment patch faint. Pharynx almost unarmed. Antenna 3 = 0-30 (0:29-0:32)
mm long, 0:20 (0:19-0:22) length of wing, 1-20 (1:17-1:26) length of 445, 1-86 (1:76-1:98) length of
labrum, one ascoid on 3-15, that on 4 about 0-39 length of segment, no papilla on 5. Wing length 1-49
(1:42-1:57) mm, 3-7 times width, R./R2:3 1°62 (1:33-1:95), R, overlap/R, 0-60 (0:51-0:69). Aedeagus
narrow and tapering, filaments about 3-8 length of pump. Paramere hooked. Style with two of spines
slightly subterminal, seta at about 0-76.
COMMENTS. The main cibarial teeth look rather like fore teeth because their long shafts point up
and therefore do not appear behind the margin of the lower wall of the cibarium. The arrange-
ment of the thick band at the bases of the teeth and of the pigment patch is like that seen in S.
perturbans.
MATERIAL EXAMINED.
West Malaysia: Gunong Besout Forest Reserve, many; for above description, 24.iii.1974-15.vi.1975
(J. Jeffery and colleagues). 10 3 measured.
DISTRIBUTION. West Malaysia: Gunong Besout area (as above); Ulu Gombak (Lewis, 1957); Gua ’Che
Yatim (Quate & Fairchild, 1961 : 220); Bukit Ibam, termite hills etc. on Fraser hill at 1360 m (L. W.
Quate, via BPBM).
Sergentomyia tracheola (Quate)
(Map 13)
Phlebotomus (Sergentomyia) tracheolus Quate, 1965 : 24. Holotype 2, PHILIPPINES (BPBM) [not examined].
305
“at9.d006 600 |
v7
Figs 243-257 Sergentomyia species. 243-250, S. pachystoma: (243) 2, head; (244) 9, cibarium;
(245, 246) 9, parts of cibarium crushed; (247) 9, pharynx; (248) 2, spermathecae; (249) 3, cibarium;
(250) 3, terminalia. 251-257, S. reidi, 2: (251) labrocibarium and pharynx; (252) cibarium; (253)
main teeth and one small tooth crushed; (254) pigment patch of same fly removed; (255) main
teeth and part of pigment patch of holotype partly flattened; (256) one tooth of Fig. 253; (257)
small teeth in fresh Berlese’s medium.
306
NO
Oo
Oo
NO
oa
N
L025
NO
Oo
Nn
Figs 258-268 Sergentomyia species. 258-263, S. reidi: (258-260) 2, sagittal section of head; (261) 9,
spermatheca; (262) 3, cibarium; (263) 3, terminalia. 264, 265, S. christophersi, 9: (264) labro-
cibarium; (265) first and second adoral sensilla. 266, 267, S. clydei, 2: maxilla, and its tip from
another angle. 268, S. eadithae, 2: tip of hypopharynx.
This is a large species with brown scutum. The cibarium of the female, with no pigment patch or
fore teeth, has eight wedge-shaped teeth and a strong arch, and the pharynx has a compact group
of teeth before the hind end. The spermatheca is pyriform.
DIsTRIBUTION. Philippines: Katanglad Mount (Quate, 1965: 18).
Subgenus SINTONIUS Nitzulescu
Sergentomyia subgenus Sintonius Nitzulescu, 1931 :273; Theodor, 1958:51; Perfil’ev, 1968 : 347.
Type-species: Phlebotomus hospitii Sinton, 1924g, by subsequent designation by Lewis, 1973a : 232.
The segmented spermatheca, usually small, characterizes this subgenus which occurs largely in
307
the drier parts of the Old World. The Oriental species exist in the west. They may be divided into
the clydei series, with 18 or fewer cibarial teeth, and the hospitii series, with 35 or more teeth and
a narrow tip to the hypopharynx. Erect abdominal hairs are discussed under S. clydei.
Sergentomyia (Sintonius) christophersi (Sinton)
(Figs 264, 265, Map 13)
Phlebotomus christophersi Sinton, 1927d:22, 24; 1927e:31; 1927f:33 [description]; 1932a: 60;
1932b : 571; 1932¢:579; 1933e: 420. Syntypes, PAKISTAN (depository unknown) [not examined].
Phlebotomus (Prophlebotomus) christophersi Sinton; Parrot, 1940 : 312.
Sergentomyia (Sintonius) christophersi (Sinton); Theodor & Mesghali, 1964 : 297 [cibarium]; Lewis,
1967 : 40; 19746 : 195; Artemiev, 1976a: 35.
Phlebotomus (Sergentomyia) christophersi Sinton; Abonnenc, 1972 : 152.
The cibarium of the female has large cornua, four or five separated teeth and a very small pig-
ment patch.
2 (extra facts). Labrum with unusual pattern of adoral and cibarial sensilla. Hypopharynx smooth.
Mandible pointed and slender. Maxilla with small teeth, seven lateral, 27 ventral and a dental depth of
0-12 mm.
In view of observations by Theodor & Mesghali on the fore teeth, a few females from several
countries were examined. None was found in those from Kenya, India, Morocco, Pakistan,
Sudan, Uganda or Yemen. There were distinct teeth in a female from Iran, and minute teeth in
one from Ethiopia.
MATERIAL EXAMINED.
Pakistan: Lahore. Others as indicated above.
DISTRIBUTION. India: Jaipur (Kaul et al., 1973 : 532); Bikaner (Sharma et al., 1973c); Delhi, Karnal,
Pinjaur, Pipli, Sahranpur (Sinton’s notes). Pakistan: Jhelum, Lahore (Lewis, 1967 : 40).
Sergentomyia (Sintonius) clydei (Sinton)
(Figs 266, 267, Map 13)
Phlebotomus clydei Sinton, 19285: 179; 1928c : 312; 1932a: 60; 1932b: 571; 1932c:579; 1933e : 420.
Lectotype 3, PAKISTAN (BMNH), designated by Lewis, 1967 : 42 [examined].
Phlebotomus (Prophlebotomus) clydei Sinton; Parrot, 1940 : 312; 1946 : 71; 1953 : 112.
Sergentomyia (Sintonius) clydei (Sinton); Theodor, 1958 : 51; Theodor & Mesghali, 1964 : 297; Lewis,
1967 : 42; Perfil’ev, 1968 : 347; Artemiev, 1976a : 37.
The female has 10-15 cibarial teeth and an almost unarmed pharynx, and the male has small
cibarial spicules arranged in groups. Among specimens from Tibi, Rajasthan, India, the cibarial
fore and hind teeth of females are large, and the length of abdominal tergite 6 of the males is
variable.
2 (extra facts). Hypopharynx with long low blunt teeth. Mandible pointed. Maxilla with three lateral
teeth, one very small, and a gap between them and the 30 ventrals, dental depth 0-09 mm.
The subgeneric character of a small number of erect hairs on abdominal tergites 2-6 is often
inapplicable to males of S. c/ydei. Sinton mentioned four to six erect hairs on segment 3 and
fewer behind. In some specimens, examined from the side and from above, there are two erect
hairs on tergite 2 and none behind. Tergite 6 normally bears no hairs but small and very small
microtrichia.
MATERIAL EXAMINED.
Pakistan: Bazid Khel, drawn.
DIsTRIBUTION. India: Keliveli, Osmanabad (V. Dhanda); Ajmer (Jaswant Singh, 1933); Jaipur, Jalor
(Kaul et al., 1973 : 533); Venkatapuram (10.vii.1966, in house on human bait, R. Reuben); Tibi (Sharma
308
et al., 19736); Bikaner (Sharma et al., 1973c); Bhavnagar, Delhi, Guntur, Junagadh, Kamptee, Karnal,
Madras, Pipli, Sahranpur (Sinton’s notes). Pakistan: Kandhkot, Karachi, Khairpur, Lahore, Mir
Muhammad, Peshawar, Rawalpindi, Tando Muhammad Khan, Taxla (Lewis, 1967 : 42); Jandola,
Khirgi (Sinton, 19285 : 179); Sargodha (Sinton’s notes).
Sinton (1932c : 71) found S. clydei widely in the plains of India.
Sergentomyia (Sintonius) eadithae (Sinton)
(Fig. 268, Map 13)
Phlebotomus eadithae Sinton, 1932c : 577 [2]; 1933c : 227; 1933e : 420 [3]; Parrot, 1940 : 312. LECTO-
TYPE 92, Inp1A (Sagar ?), labelled ‘P. 4...Type 2 Saugor... 3/8/27’ (BMNH), here designated
[examined ].
The female has about 35 large pointed cibarial teeth, in a row distinctly convex near the middle,
a large pigment patch, minute pharyngeal teeth, and a long spermatheca with about 12 segments
and a small head.
2 (extra facts). Labrum about 0:19 mm long, 0:10 length of wing. Hypopharynx smooth with narrow
forward part. Mandible rather blunt. Maxilla with 11 lateral teeth, not very wide, 24 ventrals and a dental
depth of 0-09 mm.
DISTRIBUTION. India: Sirohi (Kaul et al., 1973 : 533); Hyderabad (2 ¢, Qutubuddin, 1944 : 208); Sagar
(Madyia Pradesh, Sinton, 1932c : 577); Velur (Todupuzha, Travancore, Sinton’s notes).
Sergentomyia (Sintonius) hospitii (Sinton)
(Figs 269-273, Map 13)
Phlebotomus simillimus var. hospitii Sinton, 1924g : 261; 1927d:22, 27 [spermatheca]; 1927e : 30 [ci-
barium]; 1928c : 312. Lectotype 3, PAKISTAN (BMNH), designated by Lewis, 1967 : 44 [examined].
Phlebotomus hospitii Sinton; Sinton, 19295: 174; 1932a: 60; 1933d:420; Theodor, 19385: 172 [re-
lationship to S. thompsoni Theodor].
Sergentomyia (Sintonius) hospitii (Sinton); Lewis, 1967 : 43.
The female has 50-60 cibarial teeth, and both sexes have spines on femur 1.
2 (extra facts). Labrum with, on each side, three apical sensilla, each comprising a basal refractive cup,
a leaf-like process with narrow shoulders; two very small subapical sensilla, each having, like the apicals,
a canal through the labral cuticle; a row of narrow adoral sensilla; a few small adorals near the mid line
beneath the tip of the clypeus; and the usual round cibarial sensilla. Hypopharynx narrow anteriorly and
without teeth. Maxilla with 12 broad lateral teeth and no ventrals apart from a few vestigial specks.
MATERIAL EXAMINED.
Pakistan: Said Pur, 1 9.
DisTRIBUTION. India: Almora area, Kasauli (Sinton’s notes). Pakistan: Abbottabad, Dulai, Rawalpindi,
Said Pur (Lewis, 1967 : 44).
This species was known only from the western Himalayan foothills (Sinton, 1932a).
Sergentomyia (Sintonius) orissa Kaul & Lewis
(Map 13)
Sergentomyia (Sintonius) orissa Kaul & Lewis, 1977 : 83. Holotype 9, INDIA (BMNH) [examined].
The female has 2428 cibarial teeth, more or fewer than those of other Oriental Sintonius.
DISTRIBUTION. India: Bhubaneswar area.
309
269 a ig : 7025
273
272 27)
Figs 269-273 Sergentomyia hospitii, 2: (269) labrocibarium; (270) tip of labrum; (271) tip of hypo-
pharynx; (272) tip of mandible; (273) tip of maxilla.
Sergentomyia (Sintonius) sirohi Kaul, Dhanda & Modi
(Map 13)
Sergentomyia (Sintonius) sirohi Kaul, Dhanda & Modi, 1973 : 533. Holotype 9, INDIA (presumed in
Virus Research Centre, Poona) [not examined].
In this small species the female has 10-12 spaced pointed teeth and usually (unlike tiberiadis)
no fore teeth.
DISTRIBUTION. India: Veerwada (Kaul et al., 1973 : 535).
Sergentomyia (Sintonius) tiberiadis (Adler, Theodor & Lourie)
Phlebotomus species near clydei (Sinton); Adler & Theodor, 1929 : 284.
Phlebotomus tiberiadis Adler, Theodor & Lourie, 1930 : 537; Sinton, 1932c : 579. Syntypes, ISRAEL (TC)
[not examined ].
Sergentomyia (Sintonius) tiberiadis (Adler, Theodor & Lourie); Theodor, 1958 : 51.
The female has 10-18 cibarial teeth in comb formation, the middle ones being smaller than the
laterals.
310
Sergentomyia (Sintonius) tiberiadis pakistanica Artemiev & Saf’yanova
(Map 13)
Sergentomyia (Sintonius) tiberiadis (Adler, Theodor & Lourie); Theodor & Mesghali, 1964 : 297 [Afghani-
stan]; Lewis, 1967 : 44.
Sergentomyia (Sintonius) tiberiadis pakistanica Artemiev & Saf’yanova, 1974 : 544; Artemiev, 1976a : 38.
Syntypes 19 9, 33 g, AFGHANISTAN AND U.S.S.R. (depository unknown) [not examined].
This subspecies differs from the Ethiopian and Israel nominal subspecies in having a sperma-
theca narrowing toward the tip with more (9-12) segments, a common spermathecal duct, and
13 (10-17) cibarial teeth instead of 17-18. The authors raised the eastern form to subspecies after
considering its morphology and probable allopatricity.
2 (extra facts). Hypopharynx smooth. Maxilla with six, not very broad, lateral teeth, 27 minute ventrals,
and a dental depth of 0-10 mm.
MATERIAL EXAMINED.
Pakistan: Ahmed Khel, 1 9.
DISTRIBUTION. Pakistan: Ahmed Khel, Landi Kotal, Peshawar (Lewis, 1967 : 46).
Nomen nudum
Phlebotomus javanensis Flu, 1920 : 602; Sinton, 1928c : 322 (as javaensis); Lewis, 1973a : 254.
Flu wrote; ‘Phlebotomus papatacci komt op Java voor en is er b. v. door Professor de Meyere
beschreven onder de naam van phlebotomus javanensis’ (Phlebotomus papatacci occurs on the
island of Java and has been described as phlebotomus javanensis by Professor de Meyere). P.
papatasi is unknown in this part of the world and the name javanensis is a nomen nudum.
Aspects of biology
Much information on the biology of certain Oriental species is available in the literature, and
some of it has been summarized by Forattini (1973). The following paragraphs deal mainly with
general distribution, blood-feeding habits and seasonal distribution which is an essential con-
sideration in planning collecting expeditions.
General distribution
The limits of the Oriental Region used here are those of Delfinado & Hardy (see Lewis, 1973a)
except for the inclusion of Baltistan in Pakistan. The peripheral areas include Pakistan, Nepal,
Burma, a southern Chinese belt, Taiwan, Philippines, Mangole Island, Babar Island, Indonesia
and Sri Lanka. The subregions were defined by Wallace (1876 : maps | and 10) and Bartholomew
et al. (1911: pl. I). The north-eastern boundary was discussed by Chang (1965), and all aspects
of Indian zoogeography have been reviewed by Mani (1974). Christophers (192la; 1923) dis-
cussed mosquito distribution in India and emphasized east-west faunal changes on the lower
Indus and in the Sylhet area, a Malabar fauna with affinities in Sri Lanka and east of Calcutta,
and a relationship between Himalayan and more eastern forests.
Many species of sandflies occur mainly, not in a particular faunal region, but in parts of three
regions, which together make up a vast zone with its axis running from Morocco to India. For
ease of discussion this area is here named the Triad Zone, and its approximate boundaries
(clockwise from West Africa) are: the Atlantic, 48° N, 90° E and 10° N. Its core is the ‘Saharan’
district of Christophers (19215 : 714, Map III) and the Mediterranean Subregion of the Palae-
arctic (Christophers, 1923 : 422). The northern limit of this mainly rather dry zone approxi-
mates to the cold northern boundary of sandflies, and the eastern and southern limits to the
1000 mm isohyet.
The Oriental Region is here divided into the west (Indian and Ceylonese Subregions), centre
(Indo-Chinese and Indo-Malayan Subregions, excluding Borneo and the Philippines) and east
(Borneo and the Philippines).
ork
Table 1 and the maps show a relation between distribution and certain taxonomic groupings,
and indicate that most species occur largely in either the west, centre or east, but few in both
west and centre. Combining west with Triad, centre with Palaearctic (Manchurian) and east
with Australasian gives the following figures (percentages in brackets): west, 43 (35-8); centre,
42 (35-0), east, 35 (29-2), and west plus centre, 4 (3.2).
Table 1 The Oriental Phlebotominae showing the main area of each in the Old World. The term Triad
Zone is explained in the section on general distribution (p. 311), and many of its sandflies are widespread
in the west of the Orient. For the subgenera Parrotomyia and Neophlebotomus code letters are used to
indicate groupings mentioned in the text
Oriental
Palae-
West arctic:
Triad and Man- __— Austral-
Zone West Centre East Centre churian asian
PHLEBOTOMUS
(PHLEBOTOMUS)
papatasi oa
salehi e
(PARAPHLEBOTOMUS)
alexandri + — a mat ais ne =
nuri az
sergenti ate
(S YNPHLEBOTOMUS)
eleanorae — + = ms = =a a
(LARROUSSIUS)
betisi —
kandelakii burneyi -
keshishiani -
major major —
(ADLERIUS)
chinensis chinensis — - - = es 3 oes
longiductus + — = = = = =
(EUPHLEBOTOMUS)
argentipes = — =
kiangsuensis - — +
Dhilippinensis gouldi — — + = = a a
Dhilippinensis philippinensis - = -
(ANAPHLEBOTOMUS)
colabaensis - + = = = = =
hoepplii — — —_ < a = pa
stantoni - _ — = + = a
Ungrouped
newsteadi — +
(IDIOPHLEBOTOMUS)
asperulus _ — +
erebicolus = — —_
frondifer — - +
pholetor — —
sejunctus = — —
stellae - _ =
teshi —- a r
tubifer _ is
SERGENTOM YIA
(musai-group)
musai _ - _ ve = ce =
sah as
|
l
|
|
|
ero Alor
|
|
l
312
Table 1 (cont.)
Oriental
West
Centre
East
West
and
Centre
Palae-
arctic:
Man-
churian
Austral-
asian
(SERGENTOM YIA)
dentata
punjabensis
theodori pashtunica
(PARROTOM YIA)
A sp.
africana magna
B sp.
babu babu
babu insularis
baghdadis
barraudi
bigtii
brevicaulis
brevinervis
bukidnonis
dayapensis
denticulata
franciscana
grekovi
himalayensis
kauli
mangana
modii
palestinensis
queenslandi meridionalis
rudnicki
shorttii
spinifaucis
timorica
torrechantei
yoshimotoi
(GRASSOM YIA)
indica
(NEOPHLEBOTOMUS)
arboris
balica
Besout sp.
chakravarti
dhandai
gemmea
gombaki
hamidi
hodgsoni hodgsoni
iyengari
Jefferyi
khawi
linearis
malabarica
malayae
nankingensis
PHEHPHEGTAHRPMRRPHPHEMPAHHR PH PHROOOTR HS P
+
|
Lede
l++it¢t++0 1+
++
L+++4+4 1411
Sus
Table 1 (cont.)
Oriental
Palae-
West arctic:
Triad and Man- __ Austral-
Zone West Centre East Centre churian asian
a
|
|
|
I
perturbans qu — —
purii
quatei qu - —-
Rabok sp. un - —
Sepilok sp. un - -
silvatica qu - —
tambori ba - a
tonkinensis qu — --
traubi qu - -
zeylanica ar - -
(nicnic-group)
bailyi = 9
displicata 5 a
kachekensis = =
nicnic = =
(ungrouped)
angustipennis = =
anodontis = =
cheongi = =
dapsilidentes = =
delfinadoae = =
dentacea = =
exastis = =
Sfanglianensis = —
hassani rs =
heiseri = =
hitchensi = =
imitor = =
jamesi = =
kelantani = =
knudseni = =
lagunensis = =
losarcus = =
maai - =
mahadevani a =
montana ol 7.
morini = =
neras a oa
Okinawa sp. = =
pachystoma = =
pooi = =
reidi = =
tracheola a =
(SINTONIUS)
christophersi +
clydei + -
eadithae - +
hospitii — +
orissa = Te = =
ote
+
a
c
|
ne
|
ar tet
|
|
|
+++e+ 141
| l
l |
l l
l |
|
bb ol
|
|
l
Wi ai ee
|
|
|
|
bi+tttl 1 rte
Pb+++ 1 1 ttt ttt
l l l |
| | | ]
| l l |
ae de is 5A all las nore ee
|
|
|
|
sirohi -
tiberiadis pakistanica
Table 2 Latitude and longitude (degrees and minutes) of localities not shown in their original spelling
in the 1972 Times Atlas. All latitudes, except for Java and Nusa Tenggara, are north, and all longitudes east
Lati- Longi- Lati- Longi-
tude tude tude tude
ANDAMAN ISLANDS Nedumangad 8 36 TEO1
Chinya Tapu 11 41 92 43 Paloc 8 42 TP O2
BANGLADESH Panada-Agraharam 1632 81 35
Bongong, Tessore area, c. 23 10 89 12 Panchgani TPES 43:15
Dhurmakura, Mymensingh Patan 19 37 75 26
area = Nasirabad, c. 24 45 90 23 Pipariya, c. 23 00 73 00
Doloi Valley = Dhalai?c. 24 20 91 50 Poladpur 18 00 1s 25
Kaoraid, near Dacca, c. 23 42 90 22 Rajouri Soc 25 74 18
BORNEO: Sabah Ramanagar, Naini Tal 29°22 79 26
Gomantong rs gules Aas Wy Bo Sat | area, C.
Sepilok SoO. LT 22 Sagar, Madhya Pradesh 23.50 78 44
BURMA Sanawar, Simla area, c. 31 07 77 09
Mezali, Minbu district, c. 20 09 84 52 Singanama, c. 21 00 79 00
CHINA Soraipani, tea estate near 26 38 94 18
Fanglian, Hainan Island,c. 18 30 110 00 Mariani in Jorhat area, c.
Kachek 1919 110 22 Sukna, Darjeeling area, c. 21 O02 88 20
Kukong 24 59 =113 10 Tamilnadu = formerly — —
Lin Ko 19 54 109 43 Madras State
Masha A fae ams WI Me Tindharia 26 53 88 22
Nanking = Nan-ching 24°26 11720 Travancore = former State — —
Sainan 23 09— S17 S52 now in Kerala
Samshui county = Ho-k’ou, Undi 16235 81 28
Kwantung Zarit VUI2-52 Veerwada, Sirohi area, c. 25 53 72 58
Suilam = Siulam 22 40 113 29 Velur, Todupuzha, Travan- 9 00 77 00
Tche-Souen = Se-Tchouen = core, very approximately
Chechuan = Chih-ts’un 23-20) > 103: 30 Wadhwan = Surendranagar 22 44 71 43
Tienpao 23°20: -106:37 NEPAL
Yungan Zoo WLS Chobhar, c. 27 40 85 13
INDIA Dhunibesi, near Kathmandu, 27 42 85 19
Aligarh, Uttar Pradesh 27 54 78 04 G:
Baraga, Sagar (Shimoga) 14 07 75 00 Syabrudens, c. 28 10 85 08
area, c. NUSA TENGGARA
Borghat 18 48 13-28 Kabara, c. 10 02 120 44
Hoora, Sagar (Shimoga) 18 00 73°05 Los Palos S39" 126° 47
area, Cc. Pedang Bay 820% DhIS.25
Hosur 14 39 ipmiy, Same 906 125 48
Jog = Gersoppa 14 12 74 51 Suai Ole CS 17
Kannur, Sagar (Shimoga) 14 17 35°10 Sumba 8 47 120 24
area Tafara Cape Doze 125" Ue
Keliveli, Akola area, c. 20 40 17-05 PAKISTAN
Konehosur, Sagar (Shimoga) 14 07 75 00 Ahmed Khel, Peshawar 34 O1 71 40
area, c. area, C.
Kotelanka 16 43 81 17 Bazid Khel, Peshawar 34 Ol 71 40
Kulathurpuzha 8 54 77 03 area, Cc.
Kumsi, Sagar (Shimoga) 14 07 75 00 Bahrein 34 53 ge
area, c. Dulai 34 14 73 30
Mendhar 33 40 74 08 Gol 35 16 toe
Mohindergarh = Mahendra- 28 17 76 14 Gwadi Bee 4 76 10
garh, Hariana Khaira Gali 33 40 73 20
Munikeriti, c. 30 21 78 29 Khairpur, Khairpur district, 27 30 68 50
Naushera, Kashmir 33 10 74 12
313
Table 2 (cont)
Lati- Longi- Lati- Longi-
tude tude tude tude
Kohat area (Kohat-Hangu 33 34 7 AS Meung dist. NE. of Bangkok 15 08 102 06
valley), c. = Nakhon Rotchasima
Mir Muhammad 31 40 I4.15 area, C.
Parkuta 35 07 76 00 Pechaburi 13 06 99 57
Said Pur 33-47 72 58 Tha Li district = Ban Tha Li 17 37101 32
Saidu Sharif 33 40 72.30 VIETNAM (NORTH)
PHILIPPINES Bac Muc 22 04~=«:105 O1
Apali, c. 13.30., 4123.20 Bim Son 20 00 = 105 50
Bay, c. 14 20 =121 20 Bui Huy Tin, Bim Son 20 00 105 50
Bigti, Angat and Norza- 14 55 121 Ol area, c.
garay area, c. Cho Ganh 20 12 105 50
Cuernos de Negros 9.14. 123,10 Coc Leu 22. 28- 103.56
Dayap, c. 1410 120 20 Dong Giao 20:03:- :405:57
Jose del Monte, c. 14 48 121 12 Kep 2i23.- 406: 55
Katanglad Mountain 8 07 124 55 Lam 21 20 106 30
La Mesa dan, c. 14:50; 120.10 Le Mi 2h oo 1585
Los Arcos $239) 125° 59 Nao Phu 21-29"). 405. 20
Los Banos, c. 14 20 121 10 Pho Doan 20222 1OSei2
Malinao, c. 1330). 123520 Pho Moi 22 24 104 02
Minglanilla 10 15 123 47 Pho Oc 20 07 +105 53
Nueva Viscaya 162200 G21 20 Vin Thui 77) Of RN ik
San Francisco, Agusan 900 125 40 Yen Ley 20 30 =105 38
area, Cc. WEST MALAYSIA
San Mateo 14 42 121 08 Batang Padang 414 101 21
Silang, c. 1410 120 50 Betis 455 101 45
Sipocot, c. 13°30) 3123220 Bukit Ibam, c. 310 103 15
Valazc; 14°50. 121.07 Carey Island 3 01 101 38
Tarampitao 910 117 40 Gua ’Che Yatim, c. 430 102 10
Tungkong Manga (district) 14 48 121 12 Gunong Besout Forest 3 49 101 12
SRI LANKA Reserve
Depanama 6 51 1D ST Kuala Rompin, c. 240 103 30
Kalgoda, prob. 6 52 do od. Kuala Trengan 430 102 25
Katuwawala 6 49 719 54 Lamir, Kampong or village, 3 31 102 43
Maharagama, c. 7 32 80 04 Kuantan area
Pannipitiya O25. 27957 Lubok Paku, Kuantan areac. 3 50 103 19
TAIWAN Pulau Meranti, Kampong, 429 100 50
Tzepeng, c. 23 01 120 14 Bruas area, c.
THAILAND Rantau Panjang, c. 510-4 AOD 25
Ban Bon Dan 14°32 101,58 Tanjong Rabok 209 101 24
Doi Sutep = Ban Bon Doi_ ‘18 48 98 55 Terenggan, Pahang, c. 3 00 103 00
Sutep Ulu Gombak 319 101 46
Khou Kaen Prov. = Mwang 16 25 102 50 Ulu Langat 3 07 101 49
Khou Kaen
Meung dist. near Vien- 17 59 =102 38
tiane, c.
The western Orient
The western area has probably received many species from the Mediterranean subregion of the
Palaearctic and from the Triad Zone generally, under the influence of increasing aridity. Many
of the species belong to Phlebotomus or to the subgenera Sergentomyia or Sintonius of Sergento-
myia. P. papatasi, perhaps owing to its domestic habit, has penetrated far into the western
316
Table 3. Showing the percentages of females in five collections
INDIA WEST MALAYSIA THAILAND
Sagar Gunong'_— Bukit
(Shimoga) _Besout Igam Lamir Ban Bon
area area area village Dan
PHLEBOTOMUS
argentipes 31 — —_ 100-0 60
Philippinensis gouldi — —- -- os 20
stantoni 0:8 0:78 — -~ —
frondifer — 0-45 — —- —
SERGENTOM YIA
babu and variant 33-4 = — — —
barraudi = 7:90 1-5 — —
himalayensis or ally 2 a — —
shorttii 10-7 — — — —
arboris ES = — _ —
gemmea — 0-56 — — —
gombaki — 47:02 4-4 — —
hamidi — 0-66 21-0 —_ —
iyengari — 9-45 0-7 — _—
jefferyi — 3-78 4-4 — —
linearis 2] — — — —
malayae — 2°78 — — —
perturbans -- 5-00 —_ —
silvatica — — — — 10
tambori — 0-33 = a —_
traubi — — 53°7 _- —
bailyi 44-8 ~- — = —
cheongi — 5-89 — ~— —_
hassani — — bi | — —
knudseni — 2°33 a =
mahadevani — — — — 10
pachystoma — 0-18 Ze — —
reidi — 11-34 6°5 —— —
Totals 261 900 138 448 10
Orient, where it occurs in an area with an annual mean relative humidity of less than 70 per cent
(Sivaramakrishnaiah & Ramanathan, 1967 : 1166).
The wet area of the Western Ghats lacks some dry-adapted species but harbours some wood-
land forms known in the central Orient. The S. arboris series in India forms a reversed-‘C’ dis-
tribution pattern, with Sri Lanka in the lower arm, and is thus comparable with the Anopheles
leucosphyrus Dénitz group (Covell, 1927 : 47; Reid, 1970 : 60).
Two species comprised most of the collection (Table 3) made by H. Trapido in 1957 in the
Sagar (Shimoga) area which lies between 564 and 609 m above sea level (Work et al., 1957 : 620).
The central Orient
The centre was formerly connected to Africa by forest which persisted in north-west Indo-
Pakistan till historical times (Traub & Twisseman, 1966 : 47, 48). Theodor (19385 : 172) drew
attention to the affinity between the Oriental and Ethiopian sandfly faunas, and many species
of the central Oriental area belong to Neophlebotomus which is well represented in the Ethiopian
forest zone. This is illustrated in Table 3 which shows collections made by A. B. Knudsen, R. B.
317
Tesh and colleagues in the Gunong Besout area, and by L. W. Quate in the Bukit Ibam area
(including Kuala Rompin and Gua ’Che Yatim). There is thus a parallel with the Oriental ano-
pheline mosquitoes which were once linked by forest with the essentially primitive species of
the Afrotropical forest (Gillies & de Meillon, 1968 : 321). Similarly, the Diceromyia culicines
have evidently spread into both Afrotropical and Oriental Regions long ago and developed
separate species (Reinert, 1970).
The central area has evidently received a number of species from the Manchurian Subregion
of the Palaearctic (Lewis & Wharton, 1963: 117), which has a rich insect fauna (Gressitt,
1958 : 213) differing from that of the Mediterranean Subregion owing to the intervening inhos-
pitable Siberian Subregion. These two different Palaearctic sources and climatic differences
between the western and central Oriental areas give them rather distinctive sandfly faunas.
The central area (Table 3, columns 2 and 3), like the eastern, has a considerable number of
ungrouped species of Sergentomyia. Australasian influence is small, presumably because even
narrow seas can form a strong barrier to weakly flying sandflies, although many of these insects
can occupy land barriers between regions.
West and central areas in general
P. argentipes is related to species of the Palaearctic Manchurian Subregion and of the eastern
Orient, and it may have extended to the far west by virtue of its peridomestic habit there. Its wide
distribution may be somewhat comparable with that of the more domestic P. papatasi in the
Old World. P. argentipes has a patchy distribution in India (Smith, 1959 : 17) and in the Orient
generally. Collections by R. H. Wharton at Lamir and by D. J. Gould in Thailand (Table 3)
exemplify this and point to the need for further study. The Lamir result could have been due to
seasonal or other changes, for no P. argentipes was found during a later visit (Lewis & Killick-
Kendrick, 1973).
S. indica is closely related to all other members of the small subgenus Grassomyia which
reaches from Dakar on the Atlantic to Timor, 144° or some 16 000 km. S. indica is one of the
species which show distinctive features of structure, distribution and biology.
The environment of India (Mani, 1974 : 54) and some other countries has been greatly altered
by deforestation and agriculture. Certain areas now have few phlebotomines, and Raynal did
not find many in the lowlands of North Vietnam. West-central Taiwan (Cates & Lien, 1970 : 540)
is probably an example of the same trend.
The eastern Orient
Wallace (1876:I, 315) found the Philippines strikingly deficient in many Malayan animals,
with an approach to the Sulawesi (Celebes) fauna, and Corbet & Pendlebury (1956 : 26) em-
phasized differences between Borneo and the Philippines. Their sandfly faunas differ from each
other, apart from one species, but the Borneo sandflies are little known.
Most species of the cave-dwelling subgenus /diophlebotomus are known from the eastern
Orient, but it is represented in India and West Malaysia and probably constitutes an ancient
relict fauna.
No sandfly of the eastern area is known from Australasia.
Breeding habits
Sandfly larvae were discovered in August 1908 (Howlett, 1909 : 239), but are usually difficult or
impossible to locate. In the western Orient Napier & Smith (1926), Carter & Antonipulle (1949)
and others have collected those of P. argentipes around houses. Nowadays biological infor-
mation is being gained through laboratory colonization, and work on P. papatasi and P. argen-
tipes has been summarized by Killick-Kendrick et al. (1977).
Adult resting sites
Most sandflies are wild and shelter in natural sites such as holes in trees or among stones and
rocks, soil cracks and animal burrows. Holes in termite hills have attracted attention in Kenya
318
where they harbour the vector of ‘termite hill kala-azar’ (Lewis, 1974a: 369). In Pakistan
Lewis (1967 : 49) found sandflies in termitaries at Lahore. In India Howlett (1913) found many
sandflies in termite mounds and Modi & Dhanda (1972) reported seven species from termitaries
in Maharashtra. In the Philippines Quate (1965: 18) often found sandflies associated with
termites.
A few species are largely domestic or peridomestic, resting in dwellings by day after feeding
on man, domestic animals or geckos. P. papatasi is an important domestic species, and in Poona,
Mitra (1956) found it mainly in houses and P. argentipes mainly in stables. P. sergenti is largely
domestic. P. argentipes, in most of its range outside India and Sri Lanka, is seldom seen and
appears to be a woodland zoophilic and exophilic species. In India P. argentipes was not found
in the outdoor survey by Kaul et al. (1976), and is known to be largely peridomestic and to bite
man but to prefer cattle. It may be that in the western Orient relatively dry conditions have
driven this species into peridomestic situations.
Raynal (1936a) in Indo-China found P. stantoni rarely indoors, but S. barraudi often around
houses. Tang & Maa (1945) encountered P. hoepplii in houses.
Food of adults
Plant sugars are probably taken by most sandflies (Lewis, 1966; 1971 : 543), and the subject
was long ago brought to prominence in India (Berberian, 1966) by the effect of fruit juice on the
transmission of Leishmania donovani by P. argentipes. An extraneous cap seen on the fascicle of
an American species (Lewis, 1975a: 511) may have been acquired during plant feeding, and a
similar cap was seen on a female of S. gemmea from the Gunong Besout area. Sandflies have
been reported to feed on caterpillars (Howlett, 1909 : 240), and two females to bite a large moth
in Africa (Kirk & Lewis, 1940 : 632; Quate, 1964 : 233; W. Ruttledge, 1965, verbal communi-
cation). S. bailyi, as noted below, may bite insects.
Vertebrate blood is usually taken by females but the male of P. argentipes can suck blood from
a puncture wound (Smith, 1959 : 19) and, as noted above, has been taken with biting females
in India.
It is well known that many species of Phlebotomus bite mammals, and that at least some species
of Sergentomyia attack lizards or other cold-blooded animals, some feed from amphibia and
some from birds. Some mammal-biters feed from a range of species. Certain mammal-biters
occasionally bite lizards, and vice versa, and a few African species of Sergentomyia sometimes
attack man. One American sandfly which normally feeds on wild mammals occasionally bites
man, apparently when its normal host is absent (Lewis, 1974a : 377), and this type of behaviour
may account for some of the records of Oriental Sergentomyia biting man.
P. papatasi and P. sergenti feed on man in many parts of the Old World. George (1970) showed
that the former is primarily anthropophilic in the Lahore and Peshawar areas but can also feed
on birds, bovines, dogs and equines, and that P. sergenti can feed on birds, bovines and man.
Around Aurangabad, Dhanda & Modi (1971 : 1568) showed that most P. papatasi fed on man,
and a few on cattle, dogs and birds, and Pandyia et al. (1972) reported it biting cattle.
With regard to P. argentipes in India, Lloyd et al. (1925) reported that around Calcutta it bit
cattle mainly, and man occasionally, and Napier & Smith (1926) considered that it fed mainly
on man and cattle. Lloyd & Napier (1930) observed that P. argentipes fed almost entirely on man
and cattle. Controversy then developed (Shortt, 193la: 1047-1049; 19315; Napier, 1931) con-
cerning identification of sandflies and the reliability of blood-identification methods. P. argentipes
in the laboratory was found to be able to bite mice and rabbits (Kala-azar Commission, 1932)
and chickens (Bhattacharya et a/., 1951). It has been taken on a pony (noted under Distribution).
Dogs are unattractive to P. argentipes according to Smith (1959 : 19), and Garnham (1965 : 145)
discussed the freedom of Indian dogs from L. donovani in this connection. Dhanda & Modi
(1971 : 1568) found that P. argentipes in the Aurangabad area bit cattle mainly and man to some
extent, and Pandiya et al. (1972) reported that it bit cattle in Gujarat. Man-biting by this species
in rural India, where man and cattle are often closely associated, resembles the habits of P.
longipes Parrot & Martin in Ethiopia (Foster et al., 1972 : 440).
319
In Bengal, Das & Mukherjee (1969) considered that P. argentipes had become more zoophilic
than before. Bray (1974 : 73) suggested that either it had been diverted to cattle by irritability
to anti-malarial DDT in houses, or that it was a complex containing an anthropophilic form
which had been controlled. Sen Gupta (1975) reported that, although about 1960 laboratory-
reared sandflies would very rarely feed on man, females with human blood had recently been
found in living rooms. It may be that for some years P. argentipes had undergone a change like
that which overtook a mosquito of the Solomon Islands (Taylor, 1975 : 287).
In Sri Lanka P. argentipes occurs in cow sheds but does not seem to bite man readily (Lewis
& Killick-Kendrick, 1973).
In West Malaysia P. argentipes has been found on cattle and attacking human bait (Lewis,
1957 : 166). Lewis & Wharton (1963) reported that it fed on cows but had rarely been found
biting man.
P. kiangsuensis can bite man out of doors in West Malaysia and may perhaps attack bats
(Lewis & Wharton, 1963: 120, 124).
Tang & Maa (1945) found that P. hoeppli was attracted to human bait and probably bit man.
Raynal (1936a : 360) found that P. stantoni fed on mammals, probably rodents, and Parrot &
Clastrier (1952 : 153) suggested that it might bite man.
P. betisi bit man in a West Malaysian cave but might normally feed on bats (Lewis & Wharton,
1963 : 118, 123, 124).
Species of subgenus /diophlebotomus probably feed on bats (Lewis & Lane, 1976).
With regard to Sergentomyia, early reports of ‘P. minutus’ probably refer to S. babu or other
species, but are of some limited interest. Howlett (1913) reported that at Pusa P. minutus fed on
geckos and bit man in summer. Lloyd & Napier (1930) recorded a considerable proportion of
P. minutus biting man, but Shortt (193la) questioned the methods of identifying flies and blood
meals. Napier (1931) insisted that some ‘P. minutus’ could bite man in Calcutta, specially babu
and shorttii. Shortt (19315) commented briefly on the precipitin test, and (19326) reproduced his
193la paper. Mitra (1956 : 234) found that two species, probably S. babu and S. bailyi, bit man
readily in the laboratory.
S. punjabensis has been taken on or near animal bait without actually being seen to bite (Lewis,
1967 : 50), and Pandiya et al. (1972) reported this and a related species biting cattle.
S. babu attacks geckos in India and was said to bite man in Mauritius (Adler & Theodor,
1957), but Theodor (1965, in letter) informed me that the man-biting in Mauritius was recorded
on an old label and needed checking. S. babu can bite birds in Pakistan (George, 1970).
S. baghdadis has been found on or near animal bait in Pakistan (Lewis, 1957 : 50), and one
female examined by George (1970) had fed on man.
Raynal (1936a : 366) found that a female of S. barraudi in Tonkin had fed on man.
S. shorttii bit geckos in a cage in- India, taking up to 20 minutes or more to feed (Shortt &
Swaminath, 1931 : 544), and was said by Napier (1931) to be able to bite man.
The normal host of S. indica is unknown but, to judge from an observation on S. squami-
pleuris in Africa (Kirk & Lewis, 1951), it may well be the species seen by Howlett (1909 : 240)
to bite frogs.
The above-mentioned capture of S. gemmea in a monkey-baited trap may have been due to
attraction. S. gombaki, in the Gunong Besout area of West Malaysia, fed to repletion on the face,
hand and foot of collectors, in camp or near a tree hole, and appeared to be often attracted to
people, and in the same area S. iyengari was taken in a trap baited with leaf monkeys (Knudsen
et al., 1978). S. malayae has been found on or near animal bait, recorded by Lewis (1957 : 165)
as S. zeylanica which has bitten man in an Indian jungle (Sinton, 1928c : 320).
S. bailyi was found to have bitten fowls in Indo-China (Raynal, 1936a; 19365). A female of
this species, posted on 26.vii.1962 by the entomologist of the Department of Agriculture, Dacca,
to the Commonwealth Institute of Entomology, was stated to have come from the larva of a
grain moth, Sitotroga cerealella (Olivier). The ovaries were partly developed and the midgut
contained an inner peritrophic membrane and a large mass of cells of unknown origin, each
about 2 um long, with a large nucleus, and visible in Berlese’s medium.
Banks (1919) collected S. nicnic in one place in the Philippines where it was abundant at
320
times, was attracted to lights, attacked man persistently and gorged slowly. Quate & Rosario
(1962) questioned this account of biting but Quate (1965 : 28) reported it biting man.
S. reidi and S. traubi have been taken on or near mammal bait (Lewis, 1957 : 165). S. clydei,
known to bite mammals in Africa, has, as noted above, once been taken on human bait in India.
Seasonal prevalence
In temperate climates some species of sandflies emerge from winter diapause and pass through
one or two annual generations which determine their seasonal incidence. In the tropics seasonal
occurrence seems to depend on specific biology and local conditions. Some species occur through-
out the year. Some flourish in the dry season when breeding places are not flooded. Others are
numerous in the rains, when high humidity may favour the adults and larvae of woodland
species.
Existing knowledge of seasonal changes in the Orient is summarized below, and seems to
reflect the gradual transition from temperate to tropical conditions.
In Lahore, Pakistan, Nasir (1958) found sandflies from February to November, and George
(1970) noted a sharp decline in October and November. Dhanda & Modi (1971 : 1567, 1569)
pointed out that in Pakistan sandflies tend to vanish in winter and to appear in March and
become numerous in April and in the damp month of August, whereas in peninsular India they
persist in the milder winter in Aurangabad (Deccan) and Poona districts; around Aurangabad
sandflies were abundant throughout most of the year, were most numerous when the monsoon
began in June and diminished in December. At Pusa, in Bihar State, Howlett (1909) noted that
sandflies were common in late September and early October. Smith (1959 : 17) reported that
in north-east India sandflies might vanish in the winter in December and January, and diminish
again in midsummer, being numerous after the monsoon, from August to October; in south
India numbers were lowest in hot dry months. In Indo-China sandflies were rare in the colder
part of the winter from January to March (Raynal, 1936a : 353), and at Phnom Penh in Cam-
bodia various species seemed to disappear in winter (Parrot & Clastrier, 1952).
Around Pusa Howlett (1915) observed that larvae of P. papatasi which hatched at the start of
the cold weather pupated in late February or early March according to temperature, and Craig-
head & Das (1928) reported that the species increased somewhat in the rains. In north Bengal it
was common in April (Brunetti, 1920). In Poona district P. papatasi was common throughout
the year (Mitra, 1956), and in Bombay City seemed to diminish in the rains (Young, 1927).
Mitra (1959) reported P. sergenti from March to October in Punch and Riasi. Sinton (1924f)
encountered sandflies in Indo-Pakistan in summer, mainly in the less hot period, and found
P. major in the hills in summer.
At Gauhati in Assam adults of P. argentipes were rare from late December to early February
but larvae could be found (Kala-azar Commission, 1932 : 155). Mitra (1955 : 82) saw it through-
out the year in parts of Maharashtra State and (1956) noted that it was common all the year in
Poona district. Sinton (1924) reported that sandflies might occur all the year round in southern
and eastern India, and that P. argentipes was common and could be found in most parts of the
year all along the coast from Bengal to Sri Lanka. Brunetti (1912) recorded it throughout the
year at Calcutta, where Bhasu & Ghosh (1954a; 1955) found that it reached a peak in July and
was least numerous in January.
In China, Tang & Maa (1945) found P. hoepplii from April to July and in September and
October.
Around Pusa S. indica increased somewhat in the rains (Craighead & Das, 1928).
Raynal (1936a : 364, 366) reported that S. barraudi, an upland species in Indo-China, dis-
appeared from the northern lowlands in summer, and that in Annam S. bailyi occurred through-
out the year but diminished in the cool weather.
Natural enemies
A number of parasites and predators have been reported by Bhattacharya & Biswas (1968),
the Kala-azar Commission (1932 : 158), Lewis & Killick-Kendrick (1973), Lewis (1967), Mitra
J2k
(1956 : 234), Mitra & Mitra (1953), Mohan & Suri (1975), Napier & Smith (1926), Rathnaswamy
& Rama Krishna (1954), Shortt & Swaminath (1931) and Subramaniam & Suri (1975). Culi-
coides have been found attached to P. argentipes according to Smith & Swaminath (1932 : 183)
and Das Gupta (1964 : 6, 7).
Relation to disease
The effect of bites
P. papatasi is a very troublesome biter in parts of Pakistan, and has caused intolerable itching
which could lead to secondary infection (Howlett, 1909 : 240; Sinton, 1924 : 1036; 19255 : 702).
Viruses
The following general summary is based on the work of Tesh & Chaniotis (1975), Tesh et al.
(1975) and other authors mentioned below. Sandfly-borne viruses of vertebrates may have
evolved from arthropod viruses, and belong to three distinct serogroups, vesicular stomatitis,
phlebotomus fever and (American) Changuinola. Unlike most vertebrate arboviruses, they seem
unable to produce a significant viraemia in infected animals or man. Most individual infected
vertebrates are probably dead-end hosts, and the viruses are likely to be maintained by insect-to-
insect transovarial transmission. Vesicular stomatitis virus infects a number of animals, including
man. Chandipura virus, a member of this group, was first isolated from sick persons in India
and has been recovered from wild sandflies there (Modi & Dhanda, 1972).
The phlebotomus fever group includes at least 25 serotypes with eight in the Old World (Tesh
et al., 1977a; 1977b); most have been recovered from sandflies, and at least five cause sandfly
fever. Human involvement shows a parallel with leishmaniasis. In tropical America sporadic
cases depend on contact with sylvan sandflies, and in the Middle East and Central Asia (and in
North Africa, Perfil’ev, 1968) the vector (P. papatasi) is domestic. The view that other vectors
exist (Mattingly, 1973 : 166) was probably partly based on incomplete knowledge of the dis-
tribution of P. papatasi.
With regard to the Oriental Region, Mitra (19545) reported that sandfly fever in India was
widespread during the warm months in the Punjab plains, Delhi, Uttar Pradesh, Bihar, Bengal,
Madhya Pradesh and Bombay; in Punjab, Delhi and western Uttar Pradesh there were spring
and autumn outbreaks related to multiplication of sandflies in hot, relatively damp, weather.
Barnett (1962) isolated a number of viruses from several species of sandflies in Pakistan, and
Lewis (1967) quoted other records from that country. George (1970 : 674, 676) found plebotomus
fever in Pakistan and considered that an epidemic was likely to follow the cessation of malaria
control. Rao (1975 : 1226) recorded both Sicilian and Naples strains from India. Goverdhan
et al. (1976) isolated both of these from the Aurangabad area, mainly in June, and regarded P.
papatasi as the main vector of both forms. They demonstrated a problem previously encountered
in Pakistan, that the success of a virus survey depends largely on fluctuating prevalence of
the virus.
The sandflies from Gunong Besout Forest Reserve in West Malaysia recorded in Table 3 were
part of a collection made during a search for viruses by A. B. Knudsen and R. B. Tesh (1977,
in letter) and their colleagues. Altogether 14 636 flies (5308 of them females) were processed
but without result. Nearly all the sandflies identified were species of Sergentomyia. The survey
contributes to the world picture of phlebotomus fever virus by suggesting that flies of this genus
play no significant part in the epidemiology, despite reports of a few isolations from Sergentomyia
elsewhere. Comparable results were obtained by Tesh (1977, in letter) who processed 2500
female and male sandflies from Taiwan where, as in most of the Old World tropics, species of
Sergentomyia outnumber those of Phlebotomus.
Dermal leishmaniasis
The following summary of dermal, and that of visceral, leishmaniasis is based largely on the
work of Adler & Theodor (1957), Garnham (1973) and Napier (1946).
322
Dermal leishmaniasis, generally known as Oriental sore in the Old World, and sometimes
locally as Delhi boil or Lahore sore, is caused by the trypanosomatid protozoan parasite Leish-
mania tropica (Wright, 1903). Le. t. major Yakimov (1915) is the parasite of rural zoonotic moist
dermal leishmaniasis in the U.S.S.R., and Le. t. minor Yakimov (1915) causes the urban anthropo-
notic dry form. In the Indian subcontinent the dry form is generally prevalent (Sen Gupta, 1968).
It occurred over the western and drier parts of the Indo-Gangetic plain and was endemic in
Pakistan. It existed down the west coast of India as far as Cambay, and east as far as Delhi, and
Sinton (1925b) regarded the Bombay-Tashkent line as the eastern limit. In an epidemic in Delhi
in 1940 there were some 20000 cases (Sharma et al., 1973c). Farooq & Qutubuddin (1945)
and Sen Gupta (1967) gave further details of distribution.
P. papatasi is a well-known vector of Le. tropica in the Old World. Sinton (19245) pointed out
that it was much more widespread than the disease in Indo-Pakistan, but Sen Gupta (1967) con-
sidered that it was the principal vector. Sinton (1922 : 579; 19255: 716, 717) compared the
distribution of Oriental sore with that of P. sergenti, and Sinton & Shortt (1934) noted their
coexistence, with P. papatasi, in Karnal. Mitra (19346 : 311) considered the question of vectors
to be undecided, Sen Gupta (1968) thought P. sergenti played some part in transmission, and
Sharma et al. (1973c : 69) believed that P. sergenti might be the main vector, with P. papatasi
playing some part.
Sen Gupta (1968) mentioned that S. christophersi had been suspected as a vector, but there
seems to be no evidence for this, or even for it biting man. On the basis of this report, however,
the species has been quoted as a habitual vector in a recent paper on North Africa.
No animal reservoir was known for a long time (Mitra, 19545), and Garnham (1965 : 146)
felt that this extremely common disease had practically ceased to be a zoonosis in India, but
Sen Gupta (1968) thought one might exist. Now there is strong evidence of a gerbil reservoir
(Sharma et al., 1973b), and the probable vector among the animals (Kalra & Lewis, 1976) is
P. salehi. Mohan & Suri (1975) have isolated from it parasites which are probably Le. tropica
and may be Le. ¢. major.
House spraying for malaria control made Oriental sore rare in Pakistan (Nasir, 1964). In parts
of India spraying from 1958 onwards caused a transient disappearance for more than ten years,
but an epidemic in Bikaner reached a peak in 1971 (Sharma et al., 1973a; 1973c), and the disease
has reappeared in several areas (Kaul et al., 1976).
Visceral leishmaniasis
This disease, known as kala-azar, is caused by Le. donovani (Laveran & Mesnil, 1903), and
India is one of the few countries with no known animal reservoir, though Adler (1964) believed
the disease might have developed from a zoonosis and he and Garnham (1965) and Hoare (1955)
thought that the infection might still be found in a wild animal. The name kala-azar, meaning
black fever, came from the Garo Hills in India, where it referred to the appearance of victims.
Before treatment was discovered about 75 per cent of patients died, mostly within two years.
Sen Gupta (1968) reported a death rate of 90-95 per cent among infected people and a general
mortality of over 25 per cent in some districts. The terror of this once-deadly disease caused
many people to desert their villages (Manson-Bahr, 1946), and its severity and tendency to spread
led to enormous loss of life, depopulation and failure of agriculture (Sen Gupta, 1967).
Several epidemics occurred in Bengal and were confused with malaria till treatment failed to
prevent many deaths. Kala-azar attracted special attention when it began to invade Assam in
1875 with the development of communications (Sen Gupta, 1967a). Between then and 1917 it
swept up the Brahmaputra valley in three distinct epidemic waves. There were epidemic periods
of about ten years and inter-epidemic periods of 15-20 years (Shortt, 1945). Before 1946 the
disease was known to be widely distributed in India but the epidemic areas were well defined
(Napier, 1926 : 224). Infections occurred near Cape Cormorin and in Madras, and from there
the coast was free till the Ganges delta. The plains of Bengal were heavily infected, and the en-
demic area extended along the Ganges plain into Bihar and to the eastern side of Uttar Pradesh
as far as Lucknow. To the north-east, Assam was heavily infected as far as Sibsagar. Extensions
323
60 U T
=~
‘ i y y 2 v
ve, 130
\e8 a . oy oa i
5 ae ec) The Oriental Region showing sources ote
[30 J) @e % Po te
aS Ad ‘ of Phlebotominae Ps
e
60 T
Ae em ! T Ai
(ee ; ; 4 F : i } Re 730
. \ @ OL Known distribution in the Oriental Region of en
nee QO Ae
og Ta ames cs
S r o™
Coeer e e* Y ® Phlebotomus papatasi a ;
= 4 @
e +e . 9 w 2 salehi / cot
70 T enon T T T T T ye *130
< « . 4 ' 1 7
Wr gt Known distribution in the Oriental Region of Co
, tla =f O Phlebotomus alexandri CL) P eleanorae =
[30 vant g ea X7/ j ist i
a ore is 5 a nuri S _, betisi / “ae
~ ~ 6 e \ e sergenti kandelakii Vi og
rae” sane ei : is a 30: a)
a ye burneyi / ‘
70) 80) 90, 100,
Map 3
"60 A : T T T y RC I30
. . . . c
¥ ae Known distribution in the Oriental Region of acer
fapia e GOAN
5 Be ane Phlebotomus keshishiani @ Pargentipes is
major major .. kiangsuensis eat
chinensis o .. Philippinensis y, coe ‘a
chinensis gouldi see
longiductus * .. p. philippinensis ore |
Uy T 2 Ti
Known distribution in the
} xe Phlebotomus colabaensis
Ok) hoepplii
e stantoni
A newsteadi
~O asperulus
bee Paine
80, 100,
T y
Oriental Region of
P erebicolus
. frondifer x
-- pholetor
. sejunctus
. Stellae
teshi
tubifer
ee
Map 5
T 43
rf Sergentomyia musai
dentata
6 punjabensis
a) theodori
Binal pashtunica
~ acai.
5 = ~ — ome Ee
e ® O
Ane
SB oe Ree as
Km
rere wer est el
° 500 1000
= = variant
80, 90,
100)
a
Known distribution in the Oriental
Region of
reat ssa
x
africana magna /
Ou sp.8 /
, babu babu ¥¢
ey b. insularis oe
ey
Map 6
326
"60 v ~ T T T 7 Y wr 130
Known distribution in the Oriental Region of pee
‘
O Sergentomyia baghdadis 4 S. denticulata oO
e ae barraudi * |, franciscana es
* bigtii O ,, grekovi cot”
v. brevicaulis © ,, himalayensis Pare
Q brevinervis © ,, kauli e mca
O 2 bukidnonis L] ,, Mangana e eae p
Sy dayapensis A ,, modii & or i:
- = _A@
0 500 1000 - variant
Assy 80) 90, 100,
Map 7
Teo Li a T ra 5 T i T E Orr 130
\. “_ Known distribution in the Oriental Region of oon
4¥v 2 oe V Sergentomyia palestinensis NS. yoshimotoi 7 O°
«<3 ( PP qi meridionalis ®@ ,, indica F
by J rudnicki S_., arboris cost
shorttii Oo ,, balica =
spinifaucis x ,, Besout sp. ee
timorica 4 ,, Chakravarti 4
torrechantei + ., dhandai
\
Pa
\
\
\
\
\
a
"60 T T T
V_Sergentomyia
°
oO
EJ
Ne
ae
=~ ae
T
Known distribution in the Oriental Region of
gemmea
gombaki
hamidi
hodgsoni
hodgsoni
iyengari
——
T T
x S.jefferyi
., khawi
ge linearis /
4 ,, malabarica
Q |. malayae /
Be nankingensis
60 Uy ne q T T T T Or 730
Rete: : : ; ,
\ oe Known distribution in the Oriental Region of Ca
a a “ K <<
4 @ Sergentcmyia perturbans S. silvatica e)
puri » tambori om
quatei », traubi ; ;
Rabok sp. ., tonkinensis é
if 2 F
Sepilok sp. ». zeylanica Det
— ~~ { aes
a4 x
- ’
—T —
Known distribution in the Oriental Region of
Sergentomyia bailyi
displicata
kachekensis
nicnic
angustipennis
anodontis
cheongi
T y v
S.dapsilidentes
*
N_,, delfinadoae x
x ,, dentacea ¢
+ |, exastis
a ,, fanglianensis
O° ,, hassani
S ,. heiseri
130
x
maai
t T
60 pe T T T T T nee 130
. . . . . . Cc
ye eos _. Known distribution in the Oriental Region of Ce
Peek, Sas oe.
30 5 oe c 4 H perme ronvic hitchensi + S. mahadevani “oO
i ee ae x imitor , montana - © ;
= at ig rl jamesi », morini cote
? Y oa > eae kelantani ., neras ce
eS eRe knudseni ., Okinawa sp. oe
es ~ ‘ lagunensis AE pachystoma t53 ]
oe e* 0 losarcus _ * .. pooi mS :
T T
60 -o T T T T T ue v130
\ “_ Known distribution in the Oriental Region of 0
Fa ) 3 qe
<6 s¥ eae Sergentomyia reidi S. hospitii °
ES es r fi Je eo J rit tracheola 2 ORS sp. fs et
VS — Y: \ or christophersi ++ sirohi cee
e ee clydei -» tiberiadis Pe
? ees =
ae Cee ae es Bs eadithae pakistanica es a |
>. eadee = zg a: ont Fe
100,
Map 13
from Bengal to the east were limited by mountains. The disease could occur in periodic epidemics
or in epidemic invasions. Kala-azar was in general confined to rural areas below 610 m with an
annual relative humidity of 70 per cent or more and a mean diurnal temperature range of 11 °C
or less. In Bengal there were peaks about January and November. General distribution was
described by Omran (1961), Shortt et al. (1928) and Sen Gupta (1968) who (1958) recorded an
epidemic between 1940 and 1949.
Kala-azar was rare in the west (Gajwani et al., 1967), but has appeared in Gujarat State, where
it may have been introduced (Vaishnav et al., 1970; Munshi e¢ al., 1972), and in Maharashtra
and Rajasthan (Wattal, 1973).
The disease exists in the absence of P. argentipes in Kashmir and south China, probably as
extensions of Palaearctic kala-azar. Jacob & Kalra (1951) reported it from Kashmir where P.
chinensis, the Chinese vector, and P. kandelakii occur (Lewis, 1967), and south Chinese kala-azar
was found after the war of 1937 (Omran, 1961).
There are records of kala-azar, often isolated or imported cases, from several other Oriental
countries, in some Of which P. argentipes is strongly zoophilic (Lewis, 1974a : 372): Bangladesh
(WHO, 1968), Borneo, South Vietnam, Sumatra, Thailand, West Malaysia (American Geo-
graphical Society, 1954), Burma (Rangoon, Omran, 1961), China (Canton, Theodor, 1964; Che
Kiang Province, WHO, 1968), Pakistan lowlands (Lewis, 1967), Sri Lanka (Castellani, 1904;
Senadhira, 1967) and Taiwan (leishmaniasis, Drutz et al., 1969).
With regard to transmission, Mackie (1915) urged that sandflies should be studied as possible
vectors. Christophers (1926) pointed out that kala-azar was largely associated with alluvium,
fairly heavy rainfall and the distribution (Sinton, 1925b) of P. argentipes. A heavy flagellate
infection developed in about 25 per cent of flies fed on a kala-azar patient (Knowles et al., 1924).
Shortt et al. (1920) and Shortt (1932a) showed that the infection passed forward to the cibarium.
It was later found that flies could be infected by people with post kala-azar dermal leishmaniasis.
After nearly 40 years of investigation Smith et al. (1941) showed that many infected flies fed on
330
plant juices developed forward infections. Finally, Swaminath et al. (1942) transmitted the disease
to volunteers. P. argentipes could be found throughout the year in Bengal and Assam but was
most prevalent in and just after the monsoon. Cows seemed to attract the sandfly to dwellings
but divert it from man to some extent.
A treatment campaign in 1922 had preventive value, but residual cases made complete success
impossible. Mass treatment reduced much transmission (Shortt, 1945) but did not prevent epi-
demics recurring (Sen Gupta, 1967, 1968), and even in 1937 there were many deaths (Ansari,
1962). After the last of these big epidemics ended in 1924 periodicity became disturbed, probably by
insecticide campaigns (Theodor, 1964). In 1953 DDT began to be used in the Indian national
malarial control programme (Das et al., 1976), and declining incidence was further reduced (Sen
Gupta, 1958), but when control had ceased in Bengal, Das & Mukherjee (19692) and Sen Gupta
(1973) realized the risk of some recrudescence, and Boreham (1975 : 89) suggested that houses
should be sprayed specifically against sandflies. Tests have indicated that they are normally
susceptible to insecticides (Basu & Ghosh, 1954b; Sen, 1959; Raghavan et al., 1967). Sen Gupta
(1975) reported that kala-azar, once reduced to a rarity, had increased in some areas and spread
to new ones. He called for renewed residual spraying against both malaria vectors and sandflies,
and indicated the possible value of spraying in the vicinity of infected people. Das et al. (1976)
warned of the danger of a new outbreak. Seal (1977) considered that treatment, rather than DDT,
had previously reduced kala-azar, and he reported an alarming increase and urged the study of
direct transmission from man to man. According to available information (A. B. Chowdhury,
1978, in letter) about 70 000 cases of kala-azar, with about 4000 deaths, were encountered in
August 1977 in an area comprising the four districts of Bihar State, Muzzaffapur, Samastipur,
Sitamarhi and Vaishali, and it was estimated that 30 000 more cases might exist in other districts.
Large-scale spraying and treatment were evidently needed to prevent the cases increasing to
200 000 and to end the outbreak (Anonymous, 1977).
Acknowledgements
This work was supported by a grant from the Medical Research Council.
I am very grateful to Professors A. B. Chowdhury, P. C. C. Garnham and W. Peters for
much information about leishmaniasis; to Dr S. Das, Dr V. Dhanda, Mr N. L. Kalra, Dr G. B.
Modi and Dr Rachel Reuben for sandflies from India; to Professor G. B. Fairchild for the
opportunity of studying Dr H. Trapido’s Indian collection; to Mr W. H. Cheong, Dr A. B.
Knudsen and colleagues, Mr S. Mahadevan, Mr Abu Hassan bin Omar, Dr A. Rudnick, Dr
K. Inder Singh, Dr R. Shagwan Singh and colleagues, Dr R. B. Tesh and Dr F. C. Thompson
for sandflies from West Malaysia; to my wife, Lesley, and Dr R. Killick-Kendrick for help in
collecting in Borneo; to Dr T. van Leeuwen for lending sandflies from the ZMA, to Dr W. A.
Steffan and Dr J. A. Tenorio of the BPBM who lent specimens from several countries; to the
collectors mentioned in the lists of distribution; and to Dr L. W. Quate and the late Brigadier J.
A. Sinton who presented many type- and other specimens to the BMNH.
References
In the selection of the following works a number have been omitted, particularly some old ones on
bionomics and relation to disease.
Abonnenc, E. 1967. Les phlébotomes de I’Angola. Publgdes cult. Co. Diam. Angola no. 77 : 57-122.
1972. Les phlébotomes de la région éthiopienne (Diptera, Psychodidae). Mém. Off. Rech. sci. tech.
Outre-mer no. 55 : 1-289.
& Léger, N. 1976. Sur une classification rationelle des Diptéres Phlebotomidae. Cah. Off. Rech. sci.
tech. Outre-mer, Ent. méd. 14 : 69-78.
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Index
Invalid names are in italics. Principal references are in bold.
A sp., 223, 228, 256, 313, 326 bailyi, 230, 293, 294, 314, 317, 320, 321, 329
Adlerius, 225, 239, 312 balica, 229, 230, 266, 270, 287, 313, 327
africana, 227, 228, 256, 257, 261, 263, 280, 313, batraudi,. 220, 222,223, 227,228,259; 261; 313;
326 St S19. 320. 321..327
alexandri, 224, 235, 312, 325 Besout sp., 223, 230, 271, 272, 313, 327
Anaphlebotomus, 224, 225, 240, 247, 312 Detisi221.. 223, 225,234, 231; 312,520, 320
angustipennis, 223, 295, 314, 329 Bibio, 233
annandalei, 241 Digti;. 223, 221, 209; 201;:3ba, O20
anodontis, 231, 232, 295, 296, 314, 329 brevicaulis, 227, 228, 256, 260, 313, 327
antennata, 222, 255 brevinervis, 223, 228, 260, 313, 327
arboris, 229, 230, 269, 270, 313, 317, 327 Brumptomyia, 221
argentipes, 218, 221, 223, 225, 240, 242, 243, 244, bukidnonis, 228, 260, 270, 300, 313, 327
LOI, Id, DUD 9115 316,020) Sel y Deey 2d, burneyi, 225, 234, 238, 312, 325
330,391
asiatica, 256 campester, 294
asperulus, 223, 224, 250, 312, 326 chakravarti, 223, 229. 271. 273;.313, 327
chalami, 292
B’sp., 223, 228, 257, 313, 326 cheongi, 219, 223, 231, 293, 296, 305, 314, 317, 329
babu, 218, 227, 228, 252, 256, 257, 258, 261, 265, chinensis, 226, 239, 240, 312, 325, 330
294, 303, 313, 317, 320, 326 christophersi, 226, 307, 308, 314, 323, 330
baghdadis, 227, 228, 257, 258, 313, 320, 327 clydei, 226, 299, 307, 308, 309, 310, 314, 321, 330
341
colabaensis, 224, 225, 247, 312, 326
crypta, 270
dapsilidentes, 231, 232, 267, 296, 314
dayapensis, 223, 228, 260, 313, 327
deccanensis, 255
delfinadoae, 231, 232, 296, 297, 314, 329
demeijerei, 283, 286, 287
dentacea, 223, 231, 297, 314, 329
dentata, 227, 253, 254, 255, 256, 313, 326
denticulata, 228, 256, 262, 313, 327
dhandai, 222, 229, 230, 269, 271, 272, 273, 313,
327
displicata, 223, 230, 294, 299, 314, 329
eadithae, 226, 307, 309, 314, 330
eleanorae, 224, 237, 312, 325
erebicolus, 223, 224, 251, 312, 326
Euphlebotomus, 225, 240, 250, 312
exastis, 223, 231, 296, 297, 314, 329
fallax, 222
fanglianensis, 231, 232, 297, 314, 329
Flebotomus, 233
franciscana, 228, 262, 300, 313, 327
frondifer, 224, 251, 283, 312, 317, 326
gemmea, 229, 230, 272, 273, 274, 313, 317, 319,
320, 328
gigas, 222
glaucus, 241
gombaki, 219, 229, 269, 274, 275, 276, 313, 317,
320, 328
gouldi, 225, 242, 245, 246, 312, 317, 325
Grassomyia, 223, 226, 268, 313, 318
grekovi, 227, 228, 256, 262, 313, 327
griseus, 238
hainanensis, 277
hamidi, 222; ‘223, 229, 270, 275; 276; 291, 313,
317, 328
hassani, 221, 231, 297, 298, 299, 314, 317, 329
heiseri, 231, 232, 299, 314, 329
hibernus, 277
himalayensis, 227, 228, 256, 262, 313, 317, 327
hindustanicus, 240
. hitchensi, 231, 232, 278, 297, 299, 301, 314, 329
hivernus, 277
hodgsoni, 229, 230, 271, 277, 313, 328
hoepplii, 225, 246, 247, 312, 319, 320, 321, 326
hospitii, 226, 307, 308, 309, 310, 314, 330
Idiophlebotomus, 223, 250, 312, 318, 320, 329
imitor, 231, 232, 299, 314, 329
indica, 223, 226, 268, 26%, 313, 318, 320, 321, 327
insularis, 227, 228, 258, 313, 326
iyengari, 219, 220, 223, 229, 230, 276, 277, 278,
291,313, 317, 320, 328
jJamesi, 221, 223, 231, 298, 300, 314, 329
javanensis, 311
jefferyi, 229, 230, 270, 278, 279, 313, 317, 328
kachekensis, 223, 230, 294, 295, 314, 329
kandelakii, 225, 234, 237, 238, 312, 325, 330
kauli, 223, 227, 261, 263, 282, 313, 327
kelantani, 231, 300, 314, 329
keshishiani, 225, 234, 238, 312, 325
khawi, 229, 230, 278, 313, 328
kiangsuensis, 225, 242, 244, 245, 312, 320, 325
ine 221, 231, 232, 298, 300, 302, 314, 317,
32
kwangsiensis, 259
lagunensis, 223, 231, 301, 314, 329
Larroussius, 225, 237, 312
linearis, 229, 230, 279, 280, 281, 288, 313, 317, 328
longiductus, 221, 226, 240, 312, 325
longipes, 319
losarcus, 231, 232, 296, 299, 301, 314, 329
Lutzomyia, 221, 302
maai, 231, 232, 301, 314, 329
magna, 227, 228, 256, 257, 313, 326
mahadevani, 223, 231, 301, 302, 314, 317, 329
major, 225, 234, 238, 239, 240, 250, 312, 321, 325
malabarica, 218, 229, 230, 263, 269, 280, 281, 282,
313,326,
malayae, 219, 220, 229, 230, 269, 281, 282, 284,
292, 313, 317, 320, 328
malayensis, 277
mangana, 227, 228, 256, 263, 264, 265, 267, 287,
313, 327
marginatus, 2A1
maynei, 248
meridionalis, 227, 228, 256, 264, 313, 327
minutus, 252, 255, 256, 258, 320
modii, 223, 228, 261, 263, 280, 282, 313, 327
montana, 231, 232, 303, 314, 329
morini, 223, 232, 303, 314, 329
musal, 222, 223, 226; 253,254,312; 326
nankingensis, 223, 229, 282, 313, 328
Neophlebotomus, 222, 223, 228, 252, 269, 295,
$12.-313, 317
neras, 231, 232, 303, 314, 329
newsteadi, 223, 225, 250, 312, 326
Newsteadia, 252
nicnic, 230, 294, 295, 314, 320, 329
niger, 257, 258
nuri, 223, 224, 236, 312, 325
Okinawa sp., 223, 231, 304, 314, 329
orissa, 226, 309, 314, 330
pachystoma, 223, 231, 232, 304, 306, 314, 317, 329
pakistanica, 226, 311, 314, 330
palestinensis, 227, 228, 264, 313, 327
panamensis, 221, 302
papatasi, 221, 224, 233, 234, 312, 316, 318, 319,
321, 322, 323, 324
papatasii, 233
Paraphlebotomus, 224, 235, 312
Parrotomyia, 220, 222, 223, 227, 256, 258, 295,
299, 303, 312, 313
pashtunica, 227, 255, 256, 313, 326.
perturbans, 218, 221, 229, 230, 283, 284, 285, 286,
287, 288, 296, 305, 314, 317, 328
philippinensis, 219, 225, 242, 245, 246, 312, 317,
325
a
Phlebotomus, 221, 223, 224, 233, 240, 253, 312,
316, 317, 319, 322
pholetor, 223, 224, 251, 312, 326
pooi, 223, 232, 305, 314, 329
poonaensis, 268
punjabensis, 226, 227, 253, 254, 255, 313, 320, 326
purii, 229, 230, 285, 288, 314, 328
quatei, 229, 230, 270, 275, 285, 288, 289, 291, 314,
328
queenslandi, 227, 228, 256, 264, 313, 327
Rabok sp., 223, 230, 290, 314, 328
reidi, 221, 231, 232, 286, 305, 306, 307, 314, 317,
321, 330
Rondanomyia, 222, 269
rudnicki, 227, 228, 261, 264, 266, 313, 327
saiehi, 224, 235, 312, 323, 324
sejunctus, 223, 224, 251, 312, 326
Sepilok sp., 223, 230, 289, 290, 314, 328
sergenti, 224, 235, 236, 312, 319, 321, 323, 325
Sergentomyia, 221, 222, 223, 226, 232, 235, 252,
253, 269, 280, 286, 300, 312, 313, 316, 317,
319, 320, 322
shortii, 227, 228, 257, 265, 313, 317, 320, 327
siamensis, 259
silvatica, 229, 230, 275, 288, 289, 290, 314, 317,
328
simillimus, 309
Sintonius, 223, 226, 307, 314, 316
sirohi, 226, 310, 314, 330
siulamensis, 259
343
A >
5
smithi, 294
spinifaucis, 223, 227, 265, 313, 327
squamipleuris, 268, 320
stantoni, 225, 247, 248, 249, 250, 287, 312, 317,
319, 320, 326
stellae, 224, 251, 312, 326
sylvaticus, 290
sylvestris, 283, 286, 287, 288
Synphlebotomus, 224, 236, 312
taiwanensis, 277, 278
tambori, 229, 230, 277, 289, 291, 293, 314, 317,
328
teshi, 223, 224, 249, 252, 312, 326
thapari, 257, 258
theodori, 227, 255, 256, 313, 326
tiberiadis, 222, 226, 310, 314, 330
timorica, 228, 265, 267, 313, 327
tonkinensis, 223, 229, 291, 292, 314, 328
torrechantei, 223, 227, 266, 267, 313, 327
tracheola, 223, 231, 305, 314, 330
traubi, 222, 229, 230, 270, 292, 314, 317, 321, 328
tubifer, 223, 224, 252, 312, 326
vanella, 300
whartoni, 283, 287
yoshimotoi, 223, 228, 263, 267, 313, 327
zeylanica, 222, 229, 230, 269, 270, 275, 282, 292,
293, 299, 314, 320, 328
zeylanicus, 241
British Museum (Natural History)
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Titles to be published in Volume 37
The anomalous ant-attended mealybugs (Homoptera:
Pseudococcidae) of south-east Asia. By D. J. Williams.
A supplementary catalogue of the family-group and genus-group
names of the Coleophoridae (Lepidoptera). By K. Sattler &
W. G. Tremewan.
A catalogue of the type-specimens of the Scarabaeinae
(Scarabaeidae) and the smaller lamellicorn families (Coleoptera)
described by G. J. Arrow. By M. E. Bacchus.
A revision of the Neotropical wasp genus Jrigonopsis Perty
(Hymenoptera: Sphecidae). By C. R. Vardy.
Nine genera of fungus-feeding Phlaeothripidae (Thysanoptera)
from the Oriental Region. By J. M. Palmer and L. A. Mound.
The phlebotomine sandflies (Diptera: Psychodidae) of the Oriental
Region. By D. J. Lewis.
Type set by John Wright & Sons Ltd, Bristol and Printed by Henry Ling Ltd, Dorchester
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